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MYCOTAXON
AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION
OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS
VOLUME ve ie. TO) See eon
COMPLETE IN TWO QUARTERLY ISSUES
CONSISTING OF vi + 582 PAGES
INCLUDING FIGURES
CO-ED! TORS
Gw Ly HENNESBERA:
French Language Editor & Book Review Editor
Laboratoire de Mycologie systématique et appliquée
Université de Louvain, B-1348 Louvain-la-Neuve, Belgium
RICHARD P. KORF
English Language Editor & Managing Editor
Plant Pathology Herbarium, Cornell University
Ithaca, New York 14853, USA
SUSAN C. GRUFF
Assistant Editor & Index Editor
Plant Pathology Herbarium, Cornell University
Ithaca, New York 14853, USA
Published by
MYCOTAXON, LTD., P.O. BOX 264, ITHACA, NY (1485035 USe
Printed in the United States of America
N INTERNATIONAL JOURNAL DESIGNED TO: EXPEDITE) PUBLICATION
OF RESEARCH ON TAXONOMY & ‘NOMENCLATURE OF FUNGI & LICHENS
Volume XVI October-December 1982 NOG
CONT ENCES
A preliminary discomycete flora of Macaronesia: Part 5,
Se Ler OCIERACCAG Mure pea es elie ec ululatGrs alors inte in inte ings LINDA) M. KOHN 1
Studies on hyphomycetes from West Bengal, India,
1. Cercospora and allied genera of West Bengal, 1, 35
ll. Cercospora and allied genera of West Bengal, 2. 58
Ill. Cereospora and allied genera of West Bengal, 3.
JO-MIN YEN, A. Ke KAR AND Boo K. DAS GO
Studies on parasitic fungi from South East Asia, 45, Parasitic
Puna from Malavepag 20 nee aes sa, JO=MIN YEN AND GLORTA) LIM G6
Phytophthora operculata sp. nov.,.a new marine fungus.
Ki Ge) PGE MAND 3.1L. ALCORN 99
Thecaphora androsacina and Entyloma garllardtamum, new species
OP UStA LAG INAUeS eeu ew a Puta Mal apy earelele wae dal Mi pry KALMAN VANKY 103
A new species of Orbtlta from Canada. J..H. HAINES AND K..N, EGGER = 107
Phellinus andina Plank §& Ryv. nova sp... 5S. PLANK AND LL. RYVARDEN. 114
A reinvestigation of the north-west Himalayan Pulvinulas.
RISHI) KAUSHAL 117
Isolation and identification of Futypa armeritacae from diseased
grapevines in Washington State.
D. A. GLAWE, C. B. SKOTLAND AND Wy J.) MOLLER 123
Stilbellaceous fungi 1. DidymosttLbe.
Jv. CRANE AND’ ADRIANNA D. HEWINGS «133
Variability of spores of different basidiocarps growing on the
same mycelium in the Aphyllophorales.
ERAST! PARMASTO AND TLMI PARMASTO). 141
A new species of Endophragmiella from sclerotia of Sclerotinia
CEN s amie Sal ye wie ied PeRRLae nial Gea Gig teters braTslaie iste aie esa net MICHAEL T. DUNN. | 152
Three new species of Hypogymnta from western North America
(Lichenes: Hypogymniaceae) .
LAWRENCE H. PIKE AND MASON E. HALE, JR, 157
A new species of Parmelia (Lichenes) with protocetraric acid.
MASON EB. HALE, JR. 162
A new host for Sphaeronaemella helvellae — Pseudorhizina
PAO MOS ORE Ae La aoa 2a sg ial bie Goh icie Ener hima era 9 DONALD H. PRISTER: > 165
[CONTENTS continued overleaf]
ISSN 0093-4666 MYXNAE J6(1) 2-340 (1982)
Library of Congress Catalongue Card Number 74-7903
a Pee ae
Published quarterly by MYCOTAXON, Ltd., P.O. Box 264, Yefaba hey BASSI N
For subscription details, availability in microfilm and, migregicne ,
and availability of articles as tear sheets J hay§ (hack gover 14850
WY 04 1962
tenets =.
CONTENTS: Continued
Smistrum Longtsporum, a new Harpellales (Trichomycetes) from
chironomid guts.
MARVIN C2) WILLEAMS, ROBERT! W.)) LIGHTWARDT AND: STEPHEN W. PETERSON
Megasporoporia a new genus of resupinate polypores.
Lio RYVARDEN 4.34) BS) WRIGHT AND M. RAJCHENBERG
Smittium cellaspora, a new Harpellales (Trichomycetes) From a
CHAT ORT PHELPS e fs CA Rene Pee ain aaa ay MARVIN: Cl WILLIAMS
NOES: /on hyphomygetes :
XLIT. New species of Acrodtetys) and Pseudasptropes from
South Africa,
XELEh Concerning \Chaetopsina romantied i) ere G. MORGAN -JONES
Phycosymbiodemes in Pseudocyphellarta in New Zealand,
B, RENNER AND Di J. GALLOWAY
A note, on Sporotrtehwn gougerottl Matrouchot 1910.
MICHAEL. R, MCGINNIS: AND LIBERO AJELLO
Peratfin embedding and semithin sectioning of basidiocarp
PR SMU S 5d) ienssa win shila ataee van KELLY A. GRATTO. AND DARRYL W..GRUND
Basidzospore variation in local. popuiations of some
APA etophorales ih. sug ak ut ERAST ) PARMASTO AND TLMI PARMASTO
New Species of fungi fromthe! Yucatan ‘Peninsula. :.,. GASTON | GUZMAN
Two new families, in the Ascomycotine.
D. L. HAWKSWORTH AND M.A, SHERWOOD
Cylindvoctadium spathiohy lle sp. nove oll. : CALVIN L.. SCHOULTIES,
NABTH By EL-GHOLL ‘AND SALVATORE) A. ALFTERT, JR.
Erynia necpyraltdarum sp.nov. and Contdtobolus apiculatue,
pathogens’ ot pyralid moths: components of the misdescribed
species Entomophthora pyraitdamm [Zysomycetes: Entomoph~
POP ALES simi ira Gila heer LA MrNSENMRey Se AR MUE Mie mic ct ba he ISRAEL) BEN-ZE'EV
Survey of the Argentine Species of the Ganoderma’ Luetdum
COMPLE RRS I) avcu tah isla ret ay MARTA &. BAZZALO AND. JORGE. E, WRIGHT
Two’ mew species of Helotiales from the eastern Himalayas,
R. SHARMA AND RICHARD P.) KORE
PENG ies Lay me edits i iia rie i ie arcu an bhen Pel ott G, Li. HENNEBERT
Notices:
International Mycological Association Record of Business Meeting.
FureaerSUpeestl ons: Tor MY CONRKON: BUthORS Wire vinea vill dey weuareremre eign
[MYCOTAXON for July4September 1982 (1521-540)
was issved July 15, 1982]
167
12
183
187
192
LIAS 4
B32,
Bod
243
249
262
265
he
293
326
Sod.
535
340
MYCOTAXON
WoleexAVIS No.0, pp. 1-34 October-December 1982
A PRELIMINARY DISCOMYCETE FLORA OF MACARONESIA:
PART 5, SCLEROTINIACEAE*
LINDA M. KOHN
Department of Botany, University of Toronto
Toronto, Ontario MoS JIAl, Canada
and
Plant Pathology Herbarium, Cornell University
Ithaca, New York 14853 USA
"Extreme remedies are very appropriate for extreme diseases."
Hippocrates [tr. William Henry Rich Jones]
APHORISMS, Sect. I, 6
Order HELOTIALES
Suborder HYMENOSCYPHINEAE
Family SCLEROTINIACEAE Whetzel 1945
KEY TO THE KNOWN MACARONESIAN GENERA
ier potnecial Cupulate, “stipitate, on a «distinct. scleroti—
um with a well-differentiated rind and medulla ..2
Ps) Apothecia cupulate, stipitate, on a substratal stro-
MagmerOres stroma vlacking {but if lacking, often
Wwitheemelanized..cells sat; base: of Stipe). ....... ees 3
2(1). Sclerotial medulla enveloping suscept tis-
SCS Risrid Hog Nei aes OOo Ee i OI IORIe Ciborinia
2'(1). Sclerotial medulla free of suscept tissues.
Sclerotinia
etl"). Ascospores brown at maturity ....... Lambertella
eee scospores hyaline at maturity. 2.26008. ...-00- 4
4(3'). Ectal excipulum composed of prosenchyma-
toucs celts =bound inftgel” oi. si. ee Poculum
4'(3'). Ectal excipulum composed of short cells,
MOL DCUITU Thee OL ry ster. were oe es sie eivic wales '0 oc 5
* The parts of this flora will appear in irregular order. Reprints of indi-
vidual parts will not be available for distribution.
2
5(4'). | Ectal excipulum composed of brick-shaped cells.
Lanzia
5'(4'). Ectal excipulum composed of globose cells ..... 6
OlSr te Lignicolous; ascospores greater than
10 yam LONG cece sesccecccccccecces Ciboria
OCS Foliicolous, on herbaceous’ stems, or
on mummified fruits; ascospores less than
10mm LONG cs siats sie ere stegerererens Moellerodiscus
Some taxa are identified here by the specimen number of
a "typical representative,"' either because our collections
are too Scanty to serve as types, or becatse sitter.
are no comprehensive monographs of Sclerotiniaceous
genera (such as Ciboria, Lanzia; Moellerodiscus, and
Poculum; too often type species are themselves doubtful.
CIBORIA Fuckel 1870
Key to the known Macaronesian species
]. Asci. 4-spored, .ascospores bearing a gél %shedine
1. Ciboria sp. 254
1'. Asci 8-spored, ascospores without a gel sheath
2(1"). Ascospores (11.7-) 13.7-18.6 ‘x 5.6672
3. Ciboria sp. 1182
2*(1").- Ascospores - 8.8=11.0°> (-13.2),) x) ees
3.0-Sige AYLI Sta 2. Ciboria sp. 573
1. Ciboria sp. 254
RECENT TAXONOMIC TREATMENTS: None.
PREVIOUS MACARONESIAN RECORDS: None.
KNOWN MACARONESIAN DISTRIBUTION
CANARY ISLANDS.
Tenerife. CUP-MM 254.
SUBSTRATUM: On twig. \2 «
Notes: This sparse collection is
too meagre to constitute a type
specimen. Although the_ tissue
structure of the apothecia, com-
posed of textura globulosa Ciboria sp. 254, 8
bound in gel, suggets affinities ascospores, CUP
in Moellerodiscus, the substrate, MM 254, x 1000.
an unidentified twig, is anomalous for that foliicolous
menus. + ihere is “no trace’ of''a stroma except for some
monk ly espromented, cells. at the base of the stipe. The
asci are J+ and 4-spored. Ascospores are variable
feeonape, from irregularly ellipsoid to pyriform,. though
Most trequently they . are. “slipper-—shaped," uni- or
biguttulate, (10s5-) 9 1) .0414.05 x 3./-4-4 wm... Spore
germination on water agar is uni- or bipolar.
2. Ciboria sp. 573
RECENT TAXONOMIC TREATMENTS: None.
PREVIOUS MACARONESIAN RECORDS: None. JC
KNOWN MACARONESIAN DISTRIBUTION
CANARY ISLANDS. “50
Tenerife. CUP-MM 573.
SUBSTRATUM: On branchlet in Ciboria sp. 573, 4 as-
stream. cospores, CUP-MM 573,
x 1000.
Notes: This small lignicolous collection also shows affi-
nities in Moellerodiscus primarily on the basis of the
miceOanatomy Of the ectal~ excipulum, in this fungus
eomposed, of -textura globulosa not bound in gel. Except
for dark-walled cells at the base of the stipe, no stroma
is evident. The asci are §8-spored and the pore J+.
The ascospores are one- and two-celled, uni- or bigut-
Pulate,, ellipsoid to fusoid to lacriform, 8.8-11.0 (-13-2)
x (2.9-) 3.0-3.7 (-4.4) pm
3. Ciboria sp. 1182
RECENT TAXONOMIC TREATMENTS: None.
PREVIOUS MACARONESIAN RECORDS: None.
KNOWN MACARONESIAN DISTRIBUTION
CANARY ISLANDS.
Tenerife. CUP-MM 1182.
SUBSTRATUM: On wood.
Notes: APOTHECIA scattered, arising from wood; turbi-
eae, \butr, “concolorous; discs up to: 1.5 mm’ <in: ydiam
tapering to form a_ short 7
stipe. ECTAL EXCIPULUM up i
to 80 wm wide along flanks
and stipe, narrower at the
margin, composed of hyaline
textura globulosa arranged
in chains perpendicular to Ciboria sp. 1182, apices
the hymenial surface, cells of 2 ascl, 4 ascospores,
up to. 15oume in’ diam, “most CUP-MM 1182, x 1000.
cells under 10 wm, at mar-
gin composed of textura prismatica in chains parallel
to asci. MEDULLARY EXCIPULUM a compact layer of
hyaline textura intricata possibly bound in gel, cells
up to 2.0 pm wide; medullary zone of stipe a compact
textura “porrecta oriented “parallel toy the #stipe tases.
SUBHYMENIUM poorly differentiated. ASCI clavate-cylin-
drical,*120-178 x 9-12" am, apices, ‘thickened” (up t6 916-0
pm), pore channel walls strongly J+, 8-spored. ASCO-
SPORES hyaline, obliquely uniseriate, ellipsoid to allan-
toid, bi- or multiguttulate in youth, becoming one-sep-
tate,” (115/—) 13-7-18.6 -x 5.8-6.4 pm. PARAPHYSES icyvilige
drical, simple, septate, 1.0 ym wide, Slightly inflated
aligned pices toe7.0%im, snot exceeding. themascie ;
This description is based on a small collection. The
turbinate apothecia and also the aseptate ascospores
suggest this sparse material is also immature. No true
stroma is in evidence; microscopically, the tissue sub-
tending the base of the stipe is composed of host cells
filled with hyaline hyphae. The apothecial microanatomy
is most like that of Ciboria peckiana .(Cooke)Korf sal]
though the spores in this collection are smaller and
the apothecia (as we have them here) are much different
macroscopically. This collection differs from Poculum
fitmum. “(Pers; = “Fr. Dumont,) @lso ont wood in having
an ectal excipulum composed of globose cells.
CIBORINIA Whetzel 1945
Key to the known Macaronesian species
bss Ascospores 10.6-13.8 x 4.8-6.4 ym, biguttulate,
with a prominent gel sheath 1. Ciborinia sp. 1466
I’. Ascospores (1.8-) 4.8-8.8 x (1.8-) 2.6-3.1 (-4.8) ym,
usually eguttulate, without a gel sheath.
2s, Cw thirsuta
1. Ciborinia sp. 1466
RECENT TAXONOMIC
TREATMENTS: None.
PREVIOUS MACARONESIAN
RECORDS: None.
KNOWN MACARONESIAN
DISTRIBUTION
CANARY ISLANDS.
Hierro. CUP-MM 1466.
SUBSTRATA: On sclerotia free
in soil or attached to debris. ——
Ciborinia sp. 1466, 5 ascospores,
brown-walled, rhizoidal hair from
base of stipe, section through flank
of apothecium showing outermost
cells and tomentum hyphae, CUP-MM
1Z66556 L000.
Notes: APOTHECIA solitary from tuberoid- sclerotia; disc
O.9-1.1 mm in diam, hymenium buff, concolorous with
excipular suriace; stipe up to 5 mm. long, 0.1 mm, wide,
Beadishwiorown darkening \tOachesinut, at the base. ECTAL
EXCIPULUM 55-148 jm wide, of hyaline, thin-walled
textura globulosa originating from cells of the medullary
exc DULUM. Which) turn.ouL at. a low angle to the. sur—
iocemnand. intergrade jinto the (chains, of (elobose .cells
Peapendicular,to» the, flank). surface, ,cells .up to: 20 wm
fimciamenwith thenlargcest ycells along flanks; at,.margin
outermost cells giving rise to some short tomentum hy-
phae; ectal excipulum of stipe composed of light brown-
walled, finely superficially roughened textura porrecta,
tiessoutenmost: ycells’ at ithe: , base, of sthe) stipe giving
rise to brown-walled rhizoidal hairs. MEDULLARY EXCI-
PULUM up to 24 pm wide, narrowing toward the margin,
of hyaline, thin-walled textura intricata, cells 4.0-6.5
pm broad; stipe medulla composed of hyaline, thin-wal-
led textura intricata. SUBHYMENIUM 10.5 wm wide, a
compact zone of light brown-walled textura intricata
moundein gel, icellss 2-3 am) ‘broad. /ASCI, arising from
croziers, columnar, 91-136 x 6.5-9.0 ym, pore faintly
J+, asci containing a substance along the inside wall
turning purple in Melzer's reagent, ascus apices thick-
ened (3.5-4.5 yam thick), 8-spored. ASCOSPORES uniseri-
ate, ellipsoid to allantoid to subreniform, (10.6-) 12.0
(-13.8) x 4.8-6.4 ym, biguttulate, with a prominent
gel sheath (easily seen in Melzer's reagent). PARAPHY-
SES simple, septate, filiform, slightly enlarged atgthe
tips, 1.0 ym wide, not exceeding asci. STROMA consisting
of tuberoid sclerotia, spherical with an irregular de-
pression on dorsal surface, firmly attached to the host
on ventral surface, 0.75-1.0 mm in diam; :meduliameo:
textura oblita with very heavily gelatinized) jwall.:
enveloping host vessel elements near site of attachment
to host tissue.
The ‘subreniform ascospores with “gel sheaths “and mine
purple reaction of the inner ascus wall in” Méelzenee
reagent distinguish this small but interesting collection.
Neither sclerotial nor Botrytis anamorphs were produced
in ;cultures. on DIFCO’ Corn Meal Agar... resuliincuees
the assignment here to Ciborinia rather than Botryoti-
nia, which was our field identification.
2. Ciborinia HIRSUTA Kohn & Korf, sp. nov.
RECENT TAXONOMIC TREATMENTS: None.
PREVIOUS MACARONESIAN RECORDS: None.
Apothecia solitaria ex sclerotiis; discus rubrobrunneus,
griseo-puberulentus, juventute crateriformis, deinde (sed
non profunde) cupulatus, 1-2 mm in diam., stipite variabili-
ter ad !t. cm longitudine, ad’ 250 latitudine. Exeipuiae
ectale ex cellulis formatum hyalinis, parietibus tenuibus
praeditis, inflato-lateriformibus vel globosis, ordinatis
in catenis 3-6 cellularibus ad superficiem perpendiculari-
bus, cellulis ad 15 p latis, eis extimis pilos saepe parien-
tibus. Pili hyalini praebentes 3 formas fortasse gradus
in progressu- referentes: tomenti hyphas 1-2-cellulares
ad 30 p longas cellulis apicalibus clavatis instructas;
pilos simplices multiseptatos columnares vel acuminatos
ad 70 p longos; pilos multiseptatos columnares vel acumina-
tos ad 100 p longos, basi saepe ramosus. Asci ex crocis
exorientes, cylindrici, 56-75 x 5-6 p, pori canalis pariete
J+, apicibus incrassatis. Ascosporae uniseriatae, hyalinae,
plerumque eguttulatae, late ellipsoideae, in magnitudine
Valdissime® vVariabi les, “Cl.8-)) 478-710 (28:0), exe oe
2.6-3.1 (-4.8) 3 asci ex apotheciis sub conditionibus
naturalibus collectis segregationem ascosporarum minorum
(4.8-5.3 x 3.5-4.0 p) et majorum (5.7-7.0 x 2.6-3.1 p) in
Gationibus 4:4,.6:2, 8:0. prae—
bentes. Stroma sub conditioni-
bus naturalibus sclerotium
discretum ovale ca. i-1.5 mm
longum, elevatum sed siccitate
collapsum; cortex superficiem
et dorsalum et ventralem in-
volvens, crassitie 1-2 cellu-
larum, ex textura globulosa
formatum, parietibus brunne-—
is, eis exterioribus densius
atratis; medulla ex textura
oblita parietibus gelatinosis
ex-miparte» formata, hospitis
telae vestigia includens.
APOTHECIA solitary from \\
sclerotia, disc reddish
brown, greyish-puberulent,
goblet-shaped in youth, be-
coming shallowly cupulate,
disc 1-2 mm in diam (apo-
heclas ..produced., «in, «-vitto
larger than those collected
in nature), stipe variable
Mimeenout (Up 410..Ca.. ccm,
250m wide. ECTAL, EXCIPU-—
LUM up to 35 pm wide along
flanks, of hyaline, thin-
walled, inflated, brick-
shaped to globose cells ar-
banged.,in chains. 3). to...6
Geis long, perpendicular
to the flank surface, cells
Mp. (0, 15 jm broad, the out- Ciborinia hirsuta, 5 as-
ermost cells frequently giv- cospores, non-septate
Mga rise. to a smaller,.«.¢lo- hairs emerging from the
Doce ccliy (up to 1l0.j7m-in outermost cells of the
GpamJAgin-turnhjegiving= wise Stipe, multiseptate
Pommnait Sos HAIRS.) ofv. three hairs originating from
types possibly representing the flanks, CUP-MM
three phases of development: 22/35.8% OOO.
a one- to two-celled, hya-
line tomentum hypha up _ to
SUbjm long; with. a clavate apical .cell;. a, multiseptate,
columnar to acuminate (but blunt-tipped) hair up to
pm socoadi and 70m long; a long, ~multiseptate, ico-
lumnar to acuminate hair, often branched at the base,
8
up to 100 pm long. MEDULLARY EXCIPULUM of two zones:
the outer zone of thin-walled, hyaline textura intricata
to textura prismatica with cells somewhat inflated,
oriented parallel with the flank surface, giving rise to
the inflated cells of the ectal excipulum, cells» wosite
14 wm broad; the inner zone composed of thin-walled,
hyaline to light brown-walled, Superficially granularly
roughened textura porrecta oriented parallel to the
hymenial surface, intergrading into textura intricata a-
bove stipe, cells 4-7 ym broad. SUBHYMENIUM a compact
light brown textura intricata,) cells» 255-4. 2 immovedar
ASCI arising from croziers, cylindrical, 56-75 x 5.0-5.8
wm, pore channel walls J+, apices thickened. ASCOSPORES
uniseriate, hyaline, usually eguttulate, broadly ellip-
soid, extremely variable in size, (1.8-) 4.8-7.0 (-8.8) x
(1.8-) 2.6-3.1 (-4.8) wm; asci from apothecia collected
in the field showing segregation of small ascospores
(4.8-5.3 x% 3.5-4.0) and large ascospores (5.7-7.0 x
2.6-3.1 ym) in 4:4, .6:2, and 8:0 ratios, ‘with @farge
ascospores often the basal ones in the chain of asco-
Spores; asci from apothecia produced in vitro showing
no segregation of large and small ascospores, producing
irregular numbers of ascospores (up to 9 per ascus),
ascospores iregular in size, some appearing to abort.
PARAPHYSES filiform, simple, 0.8-1.0 ym wide, not ex-
ceeding ascis STIPE ectal »‘excipulum! Jof \hyalinewees
brown, thin-walled textura porrecta with outermost
cells somewhat inflated and giving rise to hairs as
described for the receptacle and to much. shorter, non-
septate to multiseptate, narrower hairs (25-30 x 2 um)
the contents of which stain deeply in phloxine/KOH;
medullary excipulum of thin-walled, hyaline textura
porrecta, becoming brown-walled at base of stipe, ori-
ented parallel to stipe axis, cells 3-10 pm broad. STRO-
MA in nature discrete oval sclerotia, ca. 1-1.5 mm long,
raised, but collapsing on drying, brittle; rind envelop-
ing both dorsal and ventral surfaces, one to two cells
thick, of brown-walled textura globulosa, with athe
outer walls more heavily melanized, cells 7-10 ym in
diam; medulla in two zones, the inner zone of hyaline,
thin-walled textura intricata enveloping host vessel
elements and incorporating some amorphous. material,
presumably remnants of digested host tissue, cells 4-5
ym broad; the outer zone composed of hyaline textura
oblita with shorter cells than those in the inner zone
and with »thick, gelatinized walls, up to 5: am “hiek
where two adjacent walls coalesce.
CULTURE: Ascospores of CUP-MM 2273 were germinated
timmnasse-on IDIT CO iPotatoy Dextrose Agar. Cultures jwere
Takenwearom sthemiield towCorne:l SUniversity,., where “they
WweLemitansrerred: to 9 ‘cm plastic! petri, dishes containing
DUPCOM Malt “Agar;, -DIhCO” Low: spi Mycological ‘Agar,
and. DIFCO High pH Mycological Agar, After three weeks
Of incubation at room temperature under ambient light
conditions, large (0.5-2 cm long), discrete sclerotia
were «formed on malt agar, small sclerotia on Low pH
agar, aNd no, sclerotia .On sHighwpH Gacar; all ‘cultures
produced thick mats of white mycelium bearing a Myrio-
eonlum. | microconidial” ahamorph. ini» an. “attempt to
mrauce. = iormation’ ~of a-\conidial “anamorph, cultures
wenomencupated Flt. a sorowth. “chamber. at 15 -C ‘under
Ga. 21,000" lux mixed incandescent and ‘flourescent light.
While no conidial anamorph was produced, after approxi-
mately two months apothecia developed in the cultures
on malt agar. Subsequent microanatomical study of
the sclerotia showed marked dorsiventral differentiation;
the dorsal rind is similar to that described from sclero-
ifaw round in. nature, but the. ventral rind is composed
of a loose weft of dark brown-walled prosenchyma.
ine-emedulia’ ts composed of “textura oblita with thick;
Perauinized = walis, but is “loosely interwoven "in “the
ateamadjacent to the ventral rind. The entire sclerotium
Dreduceds im. culture is (overlaid by a thick layer of
white mycelium incorporating rhomboidal crystals ‘and
sporodochia of the Myrioconium anamorph.
HOMOMY PES ReP. Kort, “eM. Kohn, N. Korf & A. Y. Ross—
man, under Vaccinium, Ribeiro Frio, Madeira, Portugal,
Ziv 19/3. (CUP=MM: 2273).
KNOWN MACARONESIAN DISTRIBUTION
MADEIRA.
Madeira. CUP-MM 2272, 2273 (holotype), 2349.
SUBSTRATA: Under Vaccinium (CUP-MM 2273, 2349); under
lily (Liliaceae) (CUP-MM 2272).
Notes: On making these collections under Vaccinium
Gideaa lily bin late “spring, our initial identification
as a species of Botryotinia awaited only development
of a Botrytis conidial anamorph in cultures made from
ascospores. Subsequent cultural studies have failed
to yield a Botrytis anamorph, or any other conidial
anamorph save the Myrioconium '"microconidial" state
ubiquitous among members of the Sclerotiniaceae. This
species is consequently accommodated in the genus Cibo-
10
rinia, where it is described as new. Future collections
of this species may establish the host/substrate relation-
Ship more clearly; unfortunately the host material in
these collections is fragmentary and well-decayed.
LAMBERTELLA von Hodhnel 1918
Key to the known Macaronesian species
1. On dead branches of Myrica faya ....2. L. myricae
PRPS OT COTTIER HOSTS 7 2 srace ere ote ho dena te oe eee Sten ann oa rane at en Z
2(1'). Ascospores finely punctate, banded.
1. L. corni-maris
2'(1'). Ascospores not ornamented, not banded ..3
S027 yi Apothecia bearing bulbous-based hairs with
SUID PAILS) By sag ee aleve el ed hte tes eve ete ole’ ots Oo. Ll ezeéy lanier
suv2! ie Apothecia’ net, bearing such “hairs ...2..) een 4
4(3'). Ascospores 13.3-16.1 x 3.7-5.9 pm, with-
out a gel sheath 4. Lambertella sp. 1734
4'(3'). . Ascospores. 8.0-9.5 x. 3.2-4.2. megwien
Aree lies iad uth. <ic ss 5. Lambertella sp. 1959
1. Lambertella corni-maris Héhn., Sitzungsber. Kaiserl.
Akad. Wiss., Math.-Naturwiss. Klz, “Abtc’ "23 127:
Ss LOO
RECENT TAXONOMIC TREATMENTS: Dumont (1971).
PREVIOUS MACARONESIAN RECORDS: None.
TYPE LCOCALIT Ye ‘Austrias
KNOWN MACARONESIAN DISTRIBUTION
CANARY ISLANDS.
La Palma. CUP-MM 671.
SUBSTRATUM: unidentified petiole.
Notes: Ascospores were yellow-brown
and germinated on water agar by
single germriubes in) 24) hvs.
L. corni-maris, 4 ascospores, CUP-
MM 671, x 1000.
ata
ae Lambertella myricae Dennis & Spooner in Dennis,
Reid and Spooner, Kew Bull. 32: 113. 1977.
RECENT “TAXONOMIC TREATMENTS: * Dennis & al. (1977).
PREVIOUS MACARONESIAN RECORDS: *Dennis & al. (1977).
KNOWN MACARONESIAN DISTRIBUTION
*AZORES.
*Terceira. *Dennis & al., 26.iii1.75 n.v.
SUBSTRATUM: Dead branch of Myrica faya Ait.
Notes: The type (only known) specimen was not avail-
Bolen stOmeus cirom Kew, owhere= it 1S) con) deposit, “~since
ij Dace already “been ‘studied and’ annotated by Dr.
Ket wamoemont at ithe New ‘York Botanical Garden. Kew
Hramnor provide scoplies Of nis notes’ for our -examination.
Detloer tne caescripconm and illustration ‘of Dennis” & ‘al.
(1977) and Dumont's (1971) classification, it would
Kevemout vac Lea vapurne Whetzel’ & Dumont. in “Dumont.
Dennis & al. note their species to be "evidently very
close" to L. viburni.
3. Lambertella cfr. zeylanica Dumont, Mem. New York
Bote Gard. 22%. 167. 1971.
RECENT TAXONOMIC TREATMENTS: Dumont (1971).
PREVIOUS MACARONESIAN RECORDS: None.
pe Pee OCAW NY: 2Cevyion ;
KNOWN MACARONESIAN
DISTRIBUTION
CANARY ISLANDS.
Tenerife. CUP-MM 410.
SUBSTRATUM: On leaf blade.
Pambertella cfr. zeylanica,
ascus with ascospores, 2 hairs
from flank, CUP-MM 410, -x
1000.
12
Notes: As in the type collection of Lambertella_ zeylani-
ca, there are no brown ascospores within. ascr iy thee
‘small collection. Having examined the holotype of L. zey-
lanica (CUP-SA 3215), we find the size of the hyaline
spores in CUP-MM 410 similar to the hyaline spores
in that type. The larger, brown, discharged ascospores
in the type collection were not seen in our collecticn
from Tenerife. The acuminate hairs borne on the apothe-
cial flanks in CUP-MM 410 have much thicker walls
than those in CUP-SA 3215 and «some «collapse “oi @ceul
lumen is evident in phloxine/KOH mounts. Our determi-
Nation can only bei provisional.
4. Lambertella sp. 1734
RECENT TAXONOMIC) TREATMENTS:
None.
PREVIOUS MACARONESIAN RECORDS:
None.
KNOWN MACARONESIAN
DISTRIBUTION
AZORES.
Sao Miguel. CUP-MM 1734.
Lambertella sp. 1734
SUBSTRATUM: On Rubus sp. in l-septate hyaline as-
spray. cospores and 3-sep-
tale, pale brown-
Notes: QAPOTHECTAMccathered: widise walled germinating
Shallowly concave in youth, at ascospore, CUP-MM
maturity plane to somewhat con- P34 ee LOCO
Vex, Up. to 1-3) mm, in diam; hy—
menium warm reddish brown, margin darker than hyme-
hium, somewhat raised, exciple concolorous; ‘stipe con=—
colorous with disc, 0.5 x 13 mm. STROMA an indetermi-
nate blackened area; no stromatic cells were observed
in sections. ECTAI EXCIPULUM up: ‘to. 35 jam wide mareng
flanks, ‘of hyaline textura angularits, to) texturamolopamics
Sa,, cells’ up tow200 jim in “diam! zone narrowine (aca
gin and there composed of occasionally inflated textura
prismatica; stipe of same tissue. types as flanks, Sywitn
fine, hyaline hairs produced at the base. MEDULLARY
EXCIPULUM up to 328 ym wide at flanks, of hyaline
textura intricata, cells, 2-3 jam wide, | Stipe of hyatine
textura porrecta ‘oriented parallel to: stipe’ axis, Jeells
2-3 ym wide. SUBHYMENIUM poorly differentiated. ASCI
3
arising from croziers, clavate, 80-85 x 7-10 'ym, ascus
pore channel strongly J+, apices thickened (2-3 pm),
8-spored. ASCOSPORES biseriate at first, then biseriate
mbover and uniseriate’ below, “finally uniSeriate and
filling most of the ascus, fusoid-ellipsoid with one
Oey pon “endse tapering, 13.3-10sl x 3./-5.9 pm, one-sep—
tate and hyaline within asci, pale brown and 3-septate
at germination. PARAPHYSES filiform, once-brn]Janched,
septate, 1.5 pm broad, not exceeding asci.
Although no brown ascospores were observed within
asci, after discharge, at germination, ascospores become
Davemmooowhe ‘and. “tnrec-seprate. ihe ectal sexctpulum
temecomposea “of. téexttra’ angularis with cells “inflated
DutnotL quite so lobose.) ‘Apothecia .‘do“nots appear to: arise
trom a stroma, though the host surface’ is blackened;
thes eeackened! areas ‘eseem: 'to be ‘a irather ‘superficial
crusteaswith no ‘stromatal ‘cells in evidence’ at the junc-
tion of stipe and host. The brown-walled ascospores
and stubciborioid “exciple suggest affinities (in, Lambertel-—
Ta though -this fungus does not fully conform’ to any
‘species circumscription in that genus.
5. Lambertella sp. 1959
RECENT TAXONOMIC TREATMENTS: None.
PREVIOUS MACARONESIAN RECORDS: None.
KNOWN MACARONESIAN DISTRIBUTION
AZORES.
Terceira. CUP-MM 1959.
MADEIRA.
Madeira. CUP-MM 2355.
SUBOLRATAz{ On capsules: of sEucalyptus spp.
Norests | AVOPHECITAY scattered, .-istipitate-cupulate,, “disc
Zamm inadiam, hymentum> umber; exciple darker ochrace—
SUomeDCOWMs Stipe -Varliapleyim lenoth; “up to’ 2emm) long,
tortuous, ochraceous brown, bearing short hairs, dark
brown at base. STROMA not observed on the host, the
Pict ee waxye .culicle’ sofiythe capsule “nupturedifonly «at
iiewesive Oly Stipe attachment; the ostipe’ base, vcomposed
of a medulla of textura intricata-textura oblita with
Gelatinized jwalls,: enveloped by an ‘ectal» excipulum
of dark brown-walled textura globulosa, may constitute
the stroma. ECTAL EXCIPULUM 60 pm wide at the flanks,
14
Lambertella sp. 1959, paraphysis, ascus
apex, 8 ascospores, section through flank
of apothecium showing outermost cells of
ectal excipulum Giving rise to halite mew. |
MM 1959 5x. 1000.
of pale brown-walled, minutely granularly roughened
textura prismatica with cells somewhat inflated (barrel-
shaped), at flanks cells arranged in chains perpendi-
cular to face of receptacle; “towards “margin exci
ofcells ‘at a’ more facutes angle’ ito) thes %suriacewecer.
up? to 26.5 x 8:0. qm, outermost ‘cells’ joivinie, iicepee
hyaline to pale brown-walled, 2-3-celled, occasionally
eranularily roughened hairs up to 35 um long) bearing
a clavate, sometimes thick-walled apical ‘cell; \maram
of dark brown-walled textura prismatica in chains
av fa low! vangle to’ the “excipulammsurtaces eels Umar
Dum broad. MEDULLARY” EXCIPULUM: “al compacta zone,
up to 85 pm wide, of superficially granularly rough-
ened, hyaline textura intricata, cells .227-5.3 ym bread.
SUBHYMENIUM 48 ym wide, composed of hyaline, granu-
larly. -roughened ttextura’ intricata, |) cells)" ).6=2: 7am
broad. ASCI arising from croziers, clavate, (60-) 70-78
x (5.8-) 7.0-8.0 ym, pore channel wall J+, apices
thickened (up to 4.2 pm), 8-spored. ASCOSPORES uniseri-
ate (occasionally partially biseriate), narrowly ellipsoid
tolyreniform tor lacryform, 6.0-9.5 x 3.2-4.2 jim; hyaline
within asci, golden brown at germination, bi- to multi-
suttulate,;, bearing: a prominent gel ‘sheath up” to Oo ais
thick= ‘/PARAPHYSES ifiiliform, simple, ‘septate, 4 "Un tome
1s
PeoaG “dt apices, not exceeding “asci ol IPE: | ectal.jexci-
pulum 3-4 cells thick, of brown-walled, brick-shaped
Foumcub¢globose bcells Sorientéd yy paralte! to ‘stipe ‘axis,
Pelle Up v1tO.0 um) brodd,, ourermost cells occasionally
Giving =Gise sto clavate, | 1-Z-celled jhairs 20-25. um.) long;
medullary excipulum of hyaline, thin-walled, superficial-
iyvecranularly roughened \textura (porrecta oriented more
Mumess. parailel’ to the stipe *axis:
CUP-MM 2355, also on capsules of Eucalyptus but collec-
ted in Madeira, is similar to CUP-MM 1959 from the
PzZeress. Upon. which this ‘description 1s baséd;. but, dif—
fers in several important respects: (1) the apothecia
Coemoiaiieru (ca. linm in, diam), less. robust, and’ grey
when dry, though warm brown when rehydrated; (2)
the “cells of the excipulum ‘are shorter, narrower, and
excepmamror the hairs’ are oriented parallel to the excipu—
lar surface; (3) the ascospores are larger (10 x 3 ym)
pee cvecularly ) biguttulate. “Since bethr collections are
small, it is questionable whether these differences fall
Witirieetic: "ranve or Variation of “one Species or serve
to separate two distinct species.
LANZIA Saccardo 1884
Key to the known Macaronesian species
ie cChRO-epoLred: on husks of Castanea. 2. feiss eiec 5 a’ o's she
Ie Lanzia sp; 137
ie. easct, G-spored;,on leaf blade. 2. Lanzia sp. 409
ie eeLanzia sp. 137
RECENT TAXONOMIC TREATMENTS: None.
PREVIOUS MACARONESIAN RECORDS: None.
KNOWN MACARONESIAN DISTRIBUTION:
CANARY ISLANDS.
Tenerife. CUP-MM 137.
SUBSTRATUM: On needles of the husks of Castanea sativa
Mill.
Notes: APOTHECIA solitary, associated with stromatized
patches on the host, deeply cupulate in youth, becoming
16
Lanzia sp. “137.7 “ascus and’ #asec—
spores’ in *“Melzer’s ireagent,” pdasa—
physis, ‘two hairs: “from “the Stipe,
three hairs from apothecial flank,
CUP-MM 137, x 1000.
(LE ESS
shallowly .cupulate at maturity; disc reddish brown,
wp». to, -O./5 mm in diam; stipe. reddish. brown, p<
toward the base, up to 1 mm long. ECTALSEXCieUauUM
composed, of two jzones:; an outer (zone (3-4) céelismaiiic.
of hyaline to light brown-walled, occasionally granular-
ly roughened, brick-shaped cells 3-6 ym broad, the
outermost cells giving rise to hyaline to light brown-
walled, septate hairs 20-40 pm long usually terminating
in a clavate cell; the inner zone 20-30 um wide, tapering
off at the margin, of hyaline to light brown-walled
brick-shaped to inflated (more or less globose) cells
which are up. to 10 ym in diam; ectaly excipuloimapoe
the stipe composed of brick-shaped, brown-walled, gran-
ularly roughened cells “oriented parallel to sthesscume
axis, cells up to 5 pm broad, outermost cells occasional—
ly giving rise to one-— to multicelled tomentaum hyphae,
cells at’ base) of ‘stipe, globose; up to tlO spmoin diade
with thick, brown walls. MEDULLARY EXCIPULUM a loose
layer ca. 60 ym wide at flanks, of hyaline to light
brown-walled textura intricata, coarsely granularly
roughened, cells up to 7 ym broad; stipe of brown-wal-
led, coarsely granularly roughened prosenchyma oriented
parallel tosthe stipe axis, graduating into 7a commac.
Ley
textura intricata at the stipe base. SUBHYMENIUM a
compact layer 25 wm wide of hyaline textura intricata,
cells 1-2 ym broad. ASCI arising from croziers, 40-60 x
6-7 pm, clavate, apices thickened (up to 3 ym), J+
with ascus pore channel wall turning pale blue (darker
Dime below, and “appearing sas “a’-dark blue: ring or as
two dots at the pore channel base), 8-spored. ASCO-
SHORES suni—~ Yor biseriate, =" hyaline; ovoid-ellipsoid,
7-9 (-10) x (2-) 3 (-4) pm, minutely biguttulate. PARA-
PHYSES “filiform or’ with “apices :slightly swollen,’ 1 jam
DLoad.ersimple vor “branched, ‘septate, "not exceeding asc,
ROMAN possibly *"imited™ 410) Ystipe Wbase; “stromatized
patches on host of doubtful relationship to apothecia.
Virontunately. thes ‘collection, upon jwhich = this, description
ioe Dasced 1s. too small, to jconstitute .a type specimen.
We sweurd shave dedicated thes new species to J. T-: Pal—
mer, whose research on chestnut-burr ‘''Rustroemia"'
Sop. eUrOpen nas. been; serithorough. the sdistinctive
thick-walled hairs, tiny biguttulate spores and granu-
larly roughened medullary excipulum preclude assign-
MeMemtOn any. Species, ipresently., described, ‘in, Lanzia.
Omgethne .assumplion.= that spores in. this collection. may
jolla eG aspeen immature enough) to ~reveal.. pigmentation,
accommodation in Lambertella was also _ considered,
although no presently recognized species of that genus
matcenes, our pcollection’ either. Cultural studies are need-
Somtouselucidate thew relationship sor the: “Stromata, ‘to, the
apotnectas it should be “sought again “in#?enerife.
2. Lanzia sp. 409
RECENT TAXONOMIC
TREATMENTS: None. \
U Lanzia sp. 409,
PREVIOUS MACARONESIAN 4-spored ascus,
RECORDS: None. 0 CUP-MM 409, x
1000.
KNOWN MACARONESIAN
DISTRIBUTION
CANARY ISLANDS.
Tenerife. CUP-MM 409.
SUBSTRATUM: On leaf blade.
Notes: APOTHECIA solitary from small stromata, disc
reddish-brown, 0.5 mm in diam, darker at margin;
18
Stipe reddish brown, 190 x 150 ym. STROMA a small;
black, disc-shaped area on the adaxial leaf surface;
rind covering the dorsal stromatal surface only, com-
posed of a single layer of globose cells with outermost
cell walls melanized; medulla poorly developed with
minimal hyphal invasion of subtending host tissues.
ECTAL EXCIPULUM “of, -three. layers: «the. outer layer gaae
6.4 ym wide along flanks, of light brown textura por-
recta) “turning outt.at, ia -~high Vamgle to “the Vexcipura.
surface, cells 1.0-1.6 wm broad; middle layer two cells
thick of (light. brown,.textura prismatica, with eeu
somewhat inflated, 3.0-5.8 wm broad, oriented parallel
to receptacle surface; inner layer 8 ym broad, of light
brown textura porrecta, cells 2.7-3.2 pm broad; margin
of dark brown-walled and somewhat interwoven textura
porrecta; stipe ‘composed, of “hyaline “textura 7 pormecra
parallel’ to the Stipe axis, “cells °2-/-3./7 sim jotocaus
MEDULLARY EXCIPULUM of loosely interwoven, — Jigs
brown-walled textura intricata, cells 2-3.5 wm _ broad.
SUBHYMENTUM poorly differentiated. ASCI “from vepeatiie
croziers, clavate, 60-70 x 3-4 wm, pore channel wall
faintly J+, apices thickened (up to 3 wpm), 4-spored.
ASCOSPORES uniseriate, hyaline, ellipsoid, 2-3-multi-
euttulate, ~$.0-8°5° x 2.2~-3.0 jam. PARAPHYSES “filitarm.,
septate, sparsely jbranched,. 1) jm. broad.. ‘not Vexceedaa
asci.
In ‘the absence of pigmented ascospores, jwhicheemay
bei due to immaturity of this collection, placementiaia
Lambertella is ruled out and this species is provisional-
ly accommodated in Lanzia. While the 4-spored asci
and complex excipular ‘structure: set this species apane
the’ collection’ upom which this’ «descriptionw is =basea
is too meagre to constitute a type specimen. I[dentifica—
tion of the host and cultural studies also await acqui-
sition of additional material.
MOELLERODISCUS P. Hennings 1902
= Ciboriopsis Dennis 1962
Key to the known Macaronesian species
1. Excipular tissues of apothecia light blue in Melzer's
reagent after pretreatment: in: 10% KOH) 2g ec eee 2
i Excipulars tyssues: -notiysoy reacting @5c.si aes ore 3
1
2(1). Ascus epiplasm purple in Melzer's reagent.
1. M. iodotingens subsp. iodotingens
2'(1). Ascus epiplasm not so reacting.
2. M. iodotingens subsp. canariensis
Olle. Ascospores 3.7-5.9 (-6.6) x 1.5 pm; apothecia
on line stromata on leaves of Hedera helix.
3. M. hederae
3'(1'). Ascospores (5.6-) 6.3-8.1 x 1.4-2.2 ym; apothecia
OM ppetioles, midribs, tear blades, Vand -mummied
stone fruits. 4. Moellerodiscus sp. 1881
1. Moellerodiscus IODOTINGENS Kohn ®& Korf, sp. nov.
subsp. IODOTINGENS
RECENT TAXONOMIC TREATMENTS: None.
PREVIOUS MACARONESIAN RECORDS: None.
Bpernecia ‘solitaria vel dispersa, )in -foldorum Jaminis’,
venis, petiolisque putrescentibus stromate infectis; recep-—-
taculum cupulatum vel plano-convexum depressionem centralem
praebens, 2-3 mm in diam., discus ochraceo-—brunneis, in
sicco brunneolescens; receptaculi parietes disco pallidius
ochraceo-brunnei, furfuracei; stipes 1-2 mm longus, saltem
ad basin atrobrunneus. Excipulum ectale 20-60 yp latum,
in gelatina inclusum, ex zonis duabus compositum, zona
interiore ex textura porrecta ad superficiem receptaculi
parallela formata, parietibus tenuibus, cellulis 2-3 yp
Pacis, .20na exteriore «x textura globulosa vel “angulari
pallide brunnea formata in catenis ad superficiem receptacu-
li parallelis ordinata, cellulis 3-8 p latis, gelatina in
substantia reagente Melzerana coerulescente post usum
10% KOH antecedentum; cellulae extimae pilos moniliformes
ad 70 yp longos hyalinos vel parietibus pallide brunneis
praeditos, interdum granulari-incrustatos, efferentes,
quibus pilis abundantissimis ad marginem ubi fere semper
parietibus brunneis praediti sunt; stipitis cellulae ecta-
les ad axin stipitis perpendiculariter extentae, in gelati-
na inclusae, pilos moniliformes parietibus brunneis praedi-
tos efferentes, cellulis parietibus atrobrunneis praeditis
ad basin stipitis repertis. Excipulum medullare in latitudi-
variabile, ex textura intricata parietibus pallide brunneis
praedita ad superficiem receptaculi plus minusve parallela,
cellulis 1-2 yp latis, plerumque’ granulari-incrustatis,
cellularum contento in substantia reagente Melzerana purpur-
eo; stipitis excipulum medullare eo receptaculi simile,
20
v
M. iodotingens subsp. iodotingens, young ascus,
with repeating crozier, CUP-MM 2091, x 1000; ascus
apex showing blueing of ascus pore channel wall
in Melzer's reagent after pretreatment in 10%, KOH,
3 “ascospores, ‘excipulum near ‘the margin, vandged
hair from lower on the receptacle, CUP-MM 1880,
x 1000.
cellulis 2—3 jy latis exceptis. Asci clavati, ex aincas
enati, 8-spori, 50-60 x 5-7 p, epiplasmate in substantia
reagente Melzerana purpureo, apice incrassato (3 wp), pori
canalis pariete dilute J+ sine usu KOH antecedente, reactio-
ne aucta cum usu 10% KOH antecedente. Ascosporae ellipsoi-
deae, hyalinae, biguttulati, 7-8.8 (-10.2) x 2-3 p, biseria-
ti, germinatione tubos germinales formantes in conformatio-—
ne quae "crux coptica’’ dicitur. Paraphyses subclavatae,
1 p latae, simplices, septatae, ascos in longitudine non
excedentes.
APOTHECIA solitary to scattered on decaying stromatized
leaf blades, veins, and petioles; receptacle cupulate
to plano-convex with a central depression, 2-3 mm
in diam; disc ochraceous brown drying to /tan; surface
of receptacle paler ochraceous brown than disc, furfura-
ceous; stipe 1-2 mm long, dark brown at least at base.
ECTAL EXCIPULUM 20-60 ym wide, bound in gel, of
two zones: the inner zone of thin-walled textura porrec-
ta turning out perpendicularly to the surface of the
receptacle, cells 2-3 pm broad; outer zone of pale brown
Z1
texturade ClObuUlOsa to “{texturay “anguldaris, “oriented ~ in
Giaviceeper pendicular (tom the “surface of the “receptacle,
cells 3-8 ym broad, gel turning blue in Melzer's reagent
following pretreatment in 10% KOH; outermost cells giving
rise to hyaline to light brown-walled moniliform hairs
Upsmron 7/0 sum “longo, Voccasionally (granularly roughened,
most abundant at the margin where hairs are almost
always brown-walled; ectal cells of stipe turning out
Deependrcular toe the “stipe axis, bound in gel; giving
rise to brown-walled moniliform hairs, dark brown-wal-
ledmrceliompresent sat, tne base sou stipe. MEDULLARY EXCl
PULUM variable in breadth, of pale brown-walled textura
iivireata Torlented, snore, or esse parallel (to receptacic
surface, cells f-2 yim broad, usually granularly rough-—
ened, seal! contents purple in’ (Melzer ‘s, reagent; medulla-—
nyeexmulum of “the stipe same as that of the recepta-—
elemre xcept «cells, 2-37 4im "Droad. “ASCI*"clavate, arising
from croziers, 8-spored, 50-00, xX) 5-7) m,. epiplasm
Purple in 7Melzer's’ “reagent, “apex thickened ‘(up to 3
ym), pore channel wall faintly J+ without KOH pretreat-
mene, reaction: enhanced ‘with pretreatment in 10% KOH.
PocOo ORES biseriate, ellipsoid, hyaline, biguttulate,
Fe0=0.07 (~10.2) bx 2.0-3.0' 4im,: germinating in a’ ‘'Coptic
Cross conficuration much later followed by production
Cimamseoingie —cerm “tube. PARAPHYSES ‘subclavate, 1.0
Mimpnoade simple, septate, not exceeding “asc in, length.
MOCO hele Kort, (L.Me ekohn, N.- Kort & °A.Y. Ross=
Mange Ol decaying, stromatized leaf blades, veins, and
iMeproles ms ecUlliavated Garden, ~Cabo ‘da. Praia, Tercéira,
Azores, Portugal, 8.iv.1978. (CUP-MM 1880) (K, TFC:
ISOTYPES.) (ISOTYPES will also be distributed in Korf
& Gruff, Discomycetes Exsiccati. )
KNOWN MACARONESIAN DISTRIBUTION
AZORES. |
Flores. CUP-MM 2070, 2091, 2146, 2188 (TFC).
Sado Miguel. CUP-MM 1739.
Terceira. CUP-MM 1880 (holotype, isotypes),
1603, 119235592002",
MADEIRA.
Madeira. CUP-MM 2271.
Subot RATA: orxdecaying, stromatized leaf blades, veins
and petioles of Myrica faya, ? Pittosporum’ sp., Hedera
helix, etc., and on herbaceous stems.
Le
Notes: The tissues blueing in Melzer's reagent (at least
after pretreatment in 10% KOH) and the _ biguttulate
ascospores distinguish this species from M. tenuistipes
(Schroet.) Dumont. The larger ascospores and abundant
moniliform hairs easily distinguish it from M. musae
Dumont. Reactions in Melzer's reagent are particularly
striking in this species: with pretreatment in 10% KOH
the ectal excipulum consistently turns light to medium
blue, in Melzer's reagent; without ‘pretreatmentagc
contents of all. tissue zones are purple-browm iieg@iae
reagent.
The, “Coptic cross” spore germination Sonemag@ as:
was consistently noted for many of the collections that
were cultured, both in this subspecies and in M. iodo-
tingens subsp. canariensis (see photograph under that
subspecies). We have no explanation for this elaborate
form of germination, usually seen within 24 hours:
Only after 48 to 72 hours (at ambient room temperatures
in our field hotel/laboratories) does further Vhypias
development ensue, with usually a single, long germtube
Observed ‘from any one ‘spore. Except for melanized
cells. around ‘thes base of the stipe; associalion@wiue
stromatized substrata is purely circumstantial, since no
stromatie tissue has been, produced.’ in] our “cures
derived from ascospores.
Ze Moellerodiscus iodotingens Kohn & Korf in Kohn
subsp. CANARIENSIS Kohn, subsp. nov.
RECENT TAXONOMIC TREATMENTS: None.
PREVIOUS MACARONESIAN RECORDS: None.
Subspeciei typicae fere omni ex parte conveniens, sed
apotheciis aurantio-brunneis, excipulo ectali zona interio-
re carente, et contento cellularum in zonis omnibus recepta-—
culi non purpurascente in substantia reagente Melzerana
differt. Huius subspeciei distributio ad Insulas Canarien-
ses limitata.
Agreeing with the description of Moellerodiscus iodotin-
ens» subsp. iodotingens, in all respects except Tommie
orange-brown color of the apothecia, lack of an inner
zone Of the, ectal, excipulum,, and -lack»of+a purplegnaams
tion, in Melzer’s’ reagent: of (cell vcontéentsin Malle zone-
Of the $receptacle..', Distribution, of ithe. .subspectcema:-
contined, to the, Canary -lslands.
23
M. iodotingens subsp. ca-
nariensis, 6 ascospores (3
separated, 3 discharged in
a group) germinating on
Water -aGar in. COptic Cross |
Contiguratiom 12° hours” “atter
being © discharged; ~ CUP—-MM
1300 mex OOO UDOLOs* (ills
Korf).
ROlOMiEE: ok.b., Kort, WC. “Denison, LeM.s. Kohn & M.A.
Sherwood, on leaf blades of Prunus lusitanica L., west
of Fuente de las Pulgas, Las Yedras, Monte de las
Mercedes, Tenerife, Canary Islands, Spain, 12.i.1976.
(CUP-—MM 545)
KNOWN MACARONESIAN DISTRIBUTION
CANARY ISLANDS.
La Palma. CUP-MM 670.
Tenerife. CUP-MM 296, 445, 446, 545 (holotype),
Cole) GhEC) 9) 00/7, 000e LISI, 1186, “1309,
1310.
SUBSTRATA: On leaf blades and petioles of Prunus lusi-
tanica L. and of undetermined hosts.
Notes:), Our field notes for spore germination in the
collection from La Palma indicate germination by a
emolei.cerm: tube in .24 hours. All other collections
for which spore germination was observed showed the
typical ‘Coptic cross'' germination shown in the _ photo-
Sraph above, just as in the typical subspecies. It
is certainly possible that we overlooked "Coptic cross"
eermination jin’ =the La... Palma’ collection, - but.‘ normal
Werm tubes are not reeorded by us for other collections
until at least 48 to 72 hours after discharge. The possi-
bility of MM-670 representing yet another taxon cannot
be ignored.
3. Moellerodiscus HEDERAE Korf & Kohn, sp. nov.
RECENT TAXONOMIC TREATMENTS: None.
PREVIOUS MACARONESIAN RECORDS: None.
24
Stroma in materia viva stroma-lineare irregulare cum apothe-
ciis associatum; cortes ex cellulis saturatae pigmentosis,
aspectu frontali epidermoideis, composita. Apothecia solita-
ria vel dispersa, in paginis foliorum et adaxialibus et ab-
axialibus reperta, breviter stipitata; receptaculum patelli-
forme, 1.25-2 mm in diam.; discus rufus, in sicco brunneo-
lescens; margo integer, elevatus, disco obscurius brunneus;
receptaculi parieties. in sicto wuiis) furfuracelsec pe.
brevissimus, denigratus. Excipulum ectale zona est 25-50 yp
lata, composita ex textura globulosa parietibus tenuibus pal-
lide brunneis praedita, cellulis 10-30 p in diam.; continuum
ad marginem ubi cellularum
parietes saturatius pigmen-
VEE tosae sunt, cellulis exti-
Gy mis processus 2-cellulares
() vel pluricellulares, fili-
formes vel clavatos, 10-20
p longos et interdum aggre-
gatos in fasciculos produ-
centibus. Excipulum medul-
lare ex stratis duobus com-
positum, zone interiore 50-
80 p lata, ex texturag n=
tricata laxe intertexta
formata, cellularum parie-
tibus pallide brunnea ad
superficiem receptaculi pa-
rallela’ formata, celtutis
3-8 p latis, grosse granu-
lari-incrustatis. Subhyme-
nium zona est mediocriter
distincta, ex catenis unco-
rum iteratorum maxima pro
parte composita. Asci cy-
lindrici ‘vel ‘*subelavata-
ex uncis enati, 8-spori,
36-44 x 2.9 wp, apice in-
crassati, pori canalis pa-
riete tenuiter J+ non nisi
cum usu 10% KOH anteceden-
te. Ascospori hyalini, an-
guste ellipsoidei, 3.7-5.9
M. hederae, ectal and
medullary excipulum, as-
cus;,, »paraphysis, | 5se@tcee
ascospores, CUP-MM 2147,
x 1000.
Zo
(-6.6) x 1.5 p, oblique uniseriati vel biseriati. Paraphy-
Ses comparate |.paucac, filiformes, \séptatae, 0.5 p, latae,
ascos longitudine non excedentes.
STROMA in fresh material an irregular line stroma asso-
ciated with apothecia; rind composed of deeply pigmented
Ccollcuewhich »areg epidermoid in, face) view...) APOTHECIA
SOulahy wlOumscattered. ,occurrings son eboth gadaxialy and
abaxial sides -of leaves, short stipitate; receptacle
patelliform, 1.25-2.0 mm in diam; disc reddish brown
divin CestOmtancaemaroin:. entire, -elevated..idarker brown
thane disc;. sides. of, receptacle. reddish brown when dry,
hurturaceous; Stipe very short, blackened. ECTAL EXCI-
BULUM A a. ezone 24.-/-50) fim, awide,, of thin-walled,, light
brown-walled textura globulosa, cells 10-30 pm in diam,
cContinuang to,wmargin) where» cell (walls are’'more deeply
Pigmemmed; outermost cells giving rise ito 2—, to ‘several-
celled filiform to; clavate: processes. 10-2Z0epim long and
occasionally grouped in fascicles. MEDULLARY EXCIPULUM
composed of two layers: the inner zone, 50-80 pm_ broad,
Ciloosely interwoven textura intricata, cell walls light
brown, cells 3-4 ym broad; outer: zone, 20-30 ym broad,
Olemetighty brown - textura, porrecta, oriented parallel: “to
the surface of the receptacle, cells 3-8 wm broad with
coarsely granular incrustations. SUBHYMENIUM a poorly
daiiienentiated m zone. | composed) ~ primarily oft chains. of
Pepcaeinge. Croziers. a), AoC! | cylindrical. ‘to tsubclavate;,
arising from croziers, 8-spored, 36-44 x 2.9 pm, ascus
apex thickened, pore channel wall weakly J+ only after
pustncatmenta, with.) 10%ieKOH..- ASCOSPORES wobliquely. > uni—
seriate or biseriate, hyaline, narrowly ellipsoid, 3.7-5.9
(-6.6) x 1.5 pm. PARAPHYSES relatively few, tilitorm,
septate. 0.5 im broad; not exceeding asci in length.
Morons Reise KOrk, UsM.. Kohn, No JNorfi& A. Y.. Ross—
Moatmemonw= leaves “of » Hedera ~ helix, “iLayes das Flores,
Flores, Azores, Portugal, 14.iv.1978. (CUP-MM 2147)
KNOWN MACARONESIAN DISTRIBUTION
AZORES:
Flores. CUP-MM 2147 (holotype).
Notes: The narrowly ellipsoid ascospores, lack of a strong
ascus pore channel ‘reaction in Melzer's reagent, and
association with a line stroma with epidermoid cells
Prmtace »View «Seleathis “species apart. fromas previously
described species of Moellerodiscus. While the granular-
26
ly roughened textura -porrecta just inside ~the * globose
cells of the ectal’ excipulum has. -been consideredy sb,
others (Dumont, 1976; Spevak and Korf, 1966) to be
an inner, layer of the ectal excipulum,, dn -thigpspeci-
the layer appears to be more continuous with the
medullary excipulum and is described here’ as an Jourer
layer of that zone. Further comments on the nature
of the stroma await cultural studies from future collec-
tions, as unfortunately attempts to culture this material
were unsuccessful: no germination was noted in 48
hours, and! single,, polar. germ tubés’tonly sone iota.
times as long. as the spore were noted after, / daya,
and the cultures failed to grow ‘further. “Another species
of this genus, M. iodotingens subsp. iodotingens, occurs
on leaves of this host under the same conditions (CUP-
MM 2091, 2146), but is readily distinguished on field
and microscopic characters.
4. Moellerodiscus sp. 1881
RECENT TAXONOMIC TREATMENTS: None.
PREVIOUS MACARONESIAN RECORDS: None.
KNOWN MACARONESIAN DISTRIBUTION
AZORES.
Terceira. CUP-MM 1881 (K) (TFC), 1882.
SUBSTRATA: On~ midribs, petioles, and leaf blades,
(CUP—-MM 1881) and stone fruit mummies (CUP-MM 1882).
Notes: APOTHECIA stipitate, arising from stromatized
petioles, midribes, leaf blades, and mummified fruits;
disc shallowly cupulate to plane, hymenium orange-brown
when revived, "rosy-vinaceous" to "flesh" (Raynor)
when. fresh, receptacle somewhat furfuraceous; _ stipe
dark. brown, tapering from. the (base, off thes; recepeacian
up to 4 mm long, thin. STROMA “mummioid,” of “indeter.
minate blackened patches of host tissue; medulla of
narrow (less than 1.0 wpm wide) hyaline prosenchyma
disrupting host epidermal and cortical tissues (host
cells displaced and/or broken down); rind composed
of brown-walled textura angularis to textura globulosa,
cells 4-10 yam in diam, covering only the dorsal surface
of the stroma. , FETAL EXCIPULUM, up to: [30 pumisiwaide
27
Moellerodiscus sp. 1881, section
through flanks showing globose ectal
excipular cells giving rise to tomen-
tum hyphae, ascus apex showing J+
reaction in Melzer's reagent after pre-
treatment in 10% KOH, 9 ascospores,
CUP-MM 1881, x 1000.
along flanks, narrowing to 16 pm at the margin; along
flanks composed of hyaline, thin-walled textura globulo—
sa, cells in chains perpendicular to the apothecial sur-
face, cells up to 15 ym in diam, outermost cells giving
rise to tomentum hyphae composed of chains of hyaline,
globose to clavate cells; margin composed of prosenchyma
parallel to the asci, giving rise to tomentum hyphae
PomtOml oats lone, made up of chains of, brick-shaped
fomeitated, salobose cells ‘and terminating with” a cla—
vate cell. MEDULLARY EXCIPULUM a loose layer up to
440 ym wide along the flanks, of hyaline, pale brown-
walled) textura intricata, cells up to, 4.0 yam broad;
aummarcin a compact layer, of téextura porrecta’ parallel
to the asci. SUBHYMENIUM poorly differentiated. ASCl
Seisige irom: repeating croziers,. cylindrical,(”'55-/2) x
4-5 wm, ascus pore channel weakly J+ only after pre-
treatment with 10% KOH, 8-spored. ASCOSPORES obliquely
maiscutate,. narrowly \(.éllipsoid, one-celled, hyaline,
P520-)),6.3-8.1 ~~ 1.4-2.2, um. -PARAPHYSES cylindrical,
Bapie.s cepiate, b.0 1m wide, notJexceeding asci. ol IPE:
ectal excipulum of light brown-walled textura prismati-
Ca,, cells. somewhat inflated, oriented parallel to the
stipe axis, cells up to 6.0 wm broad; outermost cells
ear inc rise to multicellular hairs, \up: to. 45 ym long,
weeciie ends. at -a, high. angle to .the- ‘stipe: axis, . the
apical cell clavate and sometimes umbonate, longer
hairs produced at the stipe base with dark brown,
ecanularly encrusted, walls; medullary excipulum of
Meaiine, textura intricata’ more or less parallel to the
Stipe -axis,. cells 2-4 1m broad.
28
Since some.apothecia in these collections “were
associated with mummified fruits, our preliminary gene-
ric field identification was Monilinia. Isolates from
a, “culture “derived sfrom a mass ‘of ascospores JorouwT
on ‘DIFCO Malt ,Agar, High pH. Mycological Agar, ow
pH Mycological Agar, and apples yielded no ‘conidiar
anamorph except for a Myrioconium ''microconidial state."
Although the stroma is “mummioid" (Whetzel, 1945),
the small, narrow ascospores are unlike those produced
by -specieés of Monilinia. In the. absence of (a) Monta
anamorph this fungus must be accommodated in Moellero—
discus; itS. ascospores. are unlike those ~of “Crbovias
Many species of which occur on fruits. It 4s juniortumare
that’ the, host is known only from fragments ‘of } lear,
stem “and ‘fruit which could not “be identified. (Despite
the ample material, the lack of a comprehensive mono-
graph of Moellerodiscus and some hesitancy in assign-
ing, at fo, that genus lead me to, avoid deseripinowe
asm enewesDeCCLeS: alerthica time,
POCULUM Velenovsky 1934 emend. Dumont 1972
Key to the known Macaronesian species
(In cases where no stromatal tissue is apparent, the
genus Crocicreas Fr. emend. Carpenter (Leotiaceae)
should also be considered.)
Ale Percurrently proliferating, thick-walled. darkecetae
produced..from /the- innerscellsofethe eetalre<c ome
Li) Mea cheaite oe ntiteeie Ce 4. Poculum sp. 1385
Peri NOeESCLACs EDT OSECT Lat ociy 31 crstaieinpciere’s-+ siete sel «icicles see eee ys
Zl Ascus pore channel J-, even after pre-
treatment in 10% KOH 1. 'Phialea'" calopus
2) (1).8 Ascus: pore «channel |+--\. «diate ee 3
3(2'). On cupules of Castanea sativa.
3. P. sydowianum
SNe (2a) LE WOOK Beane tot rcs eaves ie cay acotpion teh ane 2. iP., (fiemun
tf. “'PhialeaY ctr. ‘calopus, (Fr?) :( Eri)y (Ovel.®, sane
S0G.) BotssFrance 26: 234. > 1879.
= Rutstroemia calopus (Fr. : Fr.) Rehm in Ra-
benhe > Krypts-Fly) Deutsch. shoes 2oamee
Pitter 391/655 1593.
Z9
“Phialea" cfr. calopus,
Sptregularly . «thickened
ascus apices, 5 asco- m
spores, CUP-MM 1373,
x 1000.
“ ae J
Pie at +2 Pe
Oy Seneca
RECENT TAXONOMIC TREATMENTS: Carpenter (1981), Dennis
(1956), White (1941).
PREVIOUS MACARONESIAN RECORDS: None.
TYPE LOCALITY: Sweden.
KNOWN MACARONESIAN DISTRIBUTION
CANARY ISLANDS.
Gomera. CUP-MM 1373.
SUBSTRATA: On herbaceous stems.
Notes. ArOlTHECIA™ solitary ‘to “scattered from blackened
Hoste tissue.) disc sup tor 2.5. mm. “in’ diam,. hymenium o—
chraceous brown, margin darker brown, excipular sur-
paces DUlierdisc narrowing to form a short stipe, 0.25-0.5
mm long, blackened at base. STROMA poorly differentia-
reaeaitnough the suriace of the host is blackened around
Eien ctipe, base; “hyphae “penetrating the host “epidermis
of dark brown-walled prosenchyma. ECTAL EXCIPULUM
MpmtOseoe une wide, bound “in “gel, ~of- thyaline textura
gtticata, cells t-4:pm broad, giving rise to a “covering
layer” 2-4 cells deep of hyaline prosenchyma oriented
parallel to the excipular surface; at margin zone nar-
rowing, cells light brown-walled. MEDULLARY EXCIPULUM
315 wm wide at flanks, of hyaline, thin-walled textura
intricata; cells 6-11.5 ym wide. SUBHYMENIUM poorly
pinterentiated, possibly bound tin. “cellw ASCI warising
from repeating croziers, cylindrical, 85-150 x $-12.4 jm,
Secuss pore Channel J—= “even after \ pretreatment with
ied, scapices,, irregularly. thickened: sup #to (3.2: yam
thick), cytoplasm dextrinoid to purplish in Melzer's
reagent after pretreatment in 2% KOH, 8-spored. ASCO-
SPORES biseriate, becoming uniseriate, hyaline, varia-
ble, broadly ellipsoid to fusoid-ellipsoid, occasionally
flattened on one side, bi- or multiguttulate (but then
often with two, prominent, larger guttules), (9.5-)
30
13.0 (414.8) se(3.7-)) 5-7) (-6.4) am. PARABH YOR Seaet
form, 1itum')broad,.'\.with a tip. 1.5. ym .brogd, exceedim.
ascic sup, (to. 5:3 4am, . branched,.. ‘septates msl iyi aegecue
excipulum of two zones: the outer zone a compact ''cover-—
ing layer" 3-5 cells deep of thin-walled, non-gelatinized
hyaline to light brown-walled prosenchyma originating
from. cells ‘of -the)/inner.-ectal. zone,“ arranped tmormsa.
less parallel to the stipe axis, with hyphal tips occasi-
onally turning . out. to form «short, . septate =lomenuua
hyphae, at base of stipe cells dark brown-walled,
embedded ‘in brown, amorphous material; inner layer
50-80 ym wide, originating from medullary cells, of
hyaline textura oblita with thick,” gelatinous” walla.
medullary excipulum of hyaline, thin-walled, “textura
intricata, not bound in gel, incorporating many rhomboi-
dalverystals, cells up to. 8 um broad, -atspoint jonas
tachment to host the cells more compact, possibly bound
in gel, continuous with hyphae disrupting the host
epidermis and penetrating subtending host tissues.
The icregularly thickened. ascus \apices anda accu
pore channel ‘walls’ do ‘not agree with )the ‘concepiage:
this species as illustrated by Dennis (1956), and our
determination is thus ‘tentative. The taxonomic, status
ol.uPeziza “calopus» br. i: Fr.-o is strl leer, “dois cae
until a) monograph, of Poculum is completed, no. teansier
of that epithet to Poculum seems advisable (see also
Carpenter, 1981).
2: Poculum firmum (Pers. : Fr.) Dumont, Mycologia
68:2 0/0. 176.
= Rutstroemia firma (Pers. : Fr.) Karst., Bidra
Kannedom Finlands Natur Folk 19: 108. 1871.
RECENT TAXONOMIC TREATMENTS: Dennis (1956), White
(1941).
PREVIOUS MACARONESIAN RECORDS: *Berkeley (1874),
+Beltrdn Tejera (1980).
ye ee LOCAL Ys ahUurope:
KNOWN MACARONESIAN DISTRIBUTION
*CANARY ISLANDS.
La Palma.) CUPSMM, 7/67, 790.3555 0 (TEGO)
*Tenerife. *Feb. 1873, on dead wood, 3500 feet
(K: specimen lost) n.v.
ot
various stages of develop-
ment, some, producing: spér=
matia, CUP-MM 767, x 1000.
O
S Pe tirmiuns. © ascospores” al
5
SUBSTRATA: On twigs.
Noles ei bismaspecies, — common 1n Europe, (is ,as yet un=
known to us from North America or Asia.
3% Poculum sydowianum (Rehm in Sydow) Dumont,
Mycologia 68: 872. 1976.
= Rutstroemia sydowiana (Rehm in Sydow) White,
Mi Wiicydia W200. 1921.
RECENT TAXONOMIC TREATMENTS: Dennis (1956), Palmer
(1964, 1968), White (1941).
PREVIOUS MACARONESIAN RECORDS: None.
fee OCA UL Ys: Germany -
KNOWN MACARONESIAN DISTRIBUTION P. sydowianum,
GANAKY ISLANDS. ascospores, CUP
La Palma. CUP-MM 874. -MM 874, x 1000
SUBSTRATUM: On cupules of Castanea sativa Mill.
Notes: Castanea burrs should be sought in Macaronesia,
Since not only this species but three others, all of
which would have been referred to Rutstroemia in some
Ireatments. [lianzia echinophila -(Bull. £ Fr.) Korf, Lan-—
zia sp. 137 (above), and Ciboria americana Durand]
are likely to occur wherever chestnut thrives. All four
could be confused easily in the field.
4. Poculum sp. 1385
RECENT TAXONOMIC TREATMENTS: None.
PREVIOUS MACARONESIAN RECORDS: None.
o2
Poculum sp. 1385, 4 as-
cospores, ascus | fapex
showing blueing of pore
channel wall in Mel-
Ze oureagent. (section
Ol gectalsexcipulum*) with
Seta “inavearly.. 2—celled
Stage, 7018 development,
and base of multicellu-
lan seta, soptical ssection
of base of developing
seta showing broken
Wall SOL voriginal seta
and percurrent prolife-
ration of another seta,
also with apex broken
Olt, CUP-MM 1385,
Spores and ascuS apex
x 1000, others x 500.
KNOWN MACARONESIAN DISTRIBUTION
CANARY ISLANDS.
Gomera. CUP-MM 1385.
SUBSTRATUM: On dying stem of Rubus sp.
Notes: APOTHECIA solitary to scattered; receptacle deep
ly cupulate to goblet-shaped, 450 ym in diam; hymenium
buff, margin chestnut brown, exterior of receptacle
buff, margin “and upper part of cups bearing “chesrius
brown setae; cup narrowing to form a short stipe 150 wm
long, buff, clothed with chestnut brown hairs, especial-
ly». at. the, ‘base. ECTAL ~EXGIPULUM™ composed )-Olmra®
zones: a compact outer zone up to 50 pm wide of hyaline
textura oblita, to textura’ intricata “with (hyphaceetain—
walled, 2-3 wm broad and immersed in a copious gel,
cells at margin brown-walled, giving rise to setae; in-
ner zone up to 30 wm wide of hyaline. textura “perrecta
more or’ less parallel to receptacle surface, ‘cells upmo
Sy)
Deli. Droad; towards, margin inner zone. merging with
Elem OuLch azone "tO form “a7 broad “gelatinized. “zone at
iiemmeiiarein. “oF LAE. originating eirom inner cells of the
OUler ectalpexcipulum,. atetirstul—3j-celledswith a clavate
apicatercerl rand with “thin, light brown). walls; walls
appearing to thicken and darken .as hair proliferates
percurrently, ultimately becoming: ‘compound (ca: 3 jm
thick); mature setae multiseptate, tapering at both
ends, 200-500 pm long, up to 20 pm wide at the broadest
point, with scars showing internal proliferation. MEDUL-
PAR ve EXCIPULUM “of —loosely, interwoven, hyaline, thin-
walled textura intricata, ‘cells up to 51m broad. SUBHY-—
MENIUM up to 50 pm wide, of hyaline, thin-walled textu-
ra intricata possibly immersed in gel, cells 1-3 jm
DroGCwmooOCt arising trom repeating “croziers, clavate;
80-105 x 10-13 pm, apex thickened (up to 4 pm), pore
channel wall J+, 8-spored. ASCOSPORES uniseriate at
first, becoming biseriate, hyaline, ellipsoid to allantoid,
usually uniguttulate , occasionally multiguttulate,
one-celled, (12-) 15-17 (-18) _x 6-7 wpm. PARAPHYSES
paomened.s iiliform, i um broad, snot exceeding .asci:
Sigiieveetissues, aS iins apothecium, outer éectaly excipular
cells giving rise to brown-walled, multiseptate, flexuous
hairs ca. 50 pm long. STROMA extremely limited; gelati-
nized ectal excipular hyphae and non-gelatinized medul-
lary hyphae ramify through host tissues, but are noti-
cably more numerous and compact at the point of stipe
emergence we i1eom = host tissue, “where: fungal celis “and
hostecuticle are covered (byea lecalized melanized ‘crust:
Unfortunately the collection upon which this description
Lome Daced =sCONntdiliS= © lLOo™ [ew = apothecia. to wconstitute a
suitable type specimen. While some germination of asco-
Sporeomon. agar was, achieved, isolates. failed, to develop
further.
Diiere aren periapsr-a Nall sdozen: or. more species
that were formerly assigned to Rutstroemia with similar
apotnecial, structure to this species. Drs. K.P. Dumont
endeh an. KOrL have, hada. joint project. on, these species
for many years (pers. comm.), of which only one, Rut-
stroemia setulata Dearness & House) White, appears
fommrave) been’ described. Other. species have been | col-
fected in the southern US; in’ Mexico, in the neotropics
and paleotropics. Whether these can be accommodated
ite Oculum,. inom which they differ: mainly in the pre-
BoucemEOLsetac, <orby constitute §a separate genus has
mor eyet been decided:
34
SCLEROTINIA Fuckel 1870, typus conservandus
= Whetzelinia Korf & Dumont 1972 (homotypic)
One known Macaronesian species
Its Sclerotinia sclerotiorum (Lib.) de Bary, Vergl.
Morpher Biol Pilze: pre22.. 16o2.
= Sclerotinia’ libertiana Fuckel, “Jahrb. ‘Nassau
schen» Vereins:! Naturk:* (23224: 33). 097
(nom. superfl.).
= Whetzelinia sclerotiorum (Lib.) Korf & Dumont
Mycologia 64: 250. 1972.
RECENT TAXONOMIC TREATMENTS: Kohn (1979).
PREVIOUS » MACARONESIAN RECORDS: +Bensaude (1926),
tt Rosalia de Sousa Dias and Lucas (1980).
IVR ED LOCADUTY ss Belotum..
KNOWN MACARONESIAN DISTRIBUTION
PALORES:
toao, Miguel. (Pereira, nev.
+t MADEIRA.
+t Madeira. Pedrosa, December 1976, n.v.
SUBSTRATA: On cultivated beet (Bensaude, 1926) and
on Salvia splendens Ker-Gawl. (Rosdlia de Sousa Dias
and Lucas, 1980).
Notes: We, have. not), collected “this. uniquitous \pathoger
of cultivated plants, but it must surely be widespread
in Macaronesia. Neither of the reports in the literature
from the region are backed up by specimens.
MYCOTAXON
VOLeAVE Ss NO. 1 (Dp s oo- 57 October-December 1982
STUDIES ON HYPHOMYCETES FROM WEST BENGAL, INDIA, I.
CERCOSPORA AND ALLIED GENERA OF WEST BENGAL, 1l.
JO-MIN YEN
Natural History Museum
900 Exposition Boulevard
Los Angeles, California 90007, U.S.A.
A. K. KAR and B. K. DAS
Department of Botany
Presidency College
Calcutta: =- 700073, india
ABSTRACT
This study of 15 species of Cercospora and allied genera
from West Bengal (India) includes descriptions and
illustrations of ten new species and a new combination:
Cercoseptoria vignicola sp. nov., on Vigna sp.;
Phaeolsariopsis argyreiae sp. nov., on Argyreia roxburghii;
Pseudocercospora alternantherae sp. nov., on Alternanthera
sp.; P. euphorbiae-piluliferae sp. nov., on Euphorbia
ilulifera; P. polyalthiae sp. nov., on Polyalthia suberosa;
P. vignigena sp. nov., on Vigna sp.; P. vitigena sp. nov, On
Vitis sp.; Stenella canthii sp. nov., on Canthiil dedymi; S.
dioscoreicola sp. nov., on Dioscorea sp.; S. stephaniae sp.
nov., on Stephania hernandifolia; and S. myxa (Sydow in
Sydow & Mitter) comb. nov., on Cordia myxa. Sipe
1. Cercoseptoria vignicola Yen, Kar & Das, sp. nov. (Fig. 1)
Maculis indistinctis. Caespitulis hypophyllis, invisibilis.
Mycelium primarium immersum: hyphis pallidissime olivaceis,
septatis, ramosis, 2-3.5ym latis. Stomatibus aliquantum
atro-brunneis, globosis 20-40um diam. Conidiophoris
primariis brevis, dense vel valde dense fasciculatis, ex
stromatibus oriundis, simplicibus, flexuosis vel undulatis,
O-l1 septatis, O-2 geniculatis, ad apicem irregulariter
rOtuUndatisS, Cicatricibus conidialis indistinctis, 22-75 x
3-5um. Mycelium secundarium superficiale: hyphis ex
stromatibus oriundis, repentis, pallide olivaceis, septatis,
ramosis, 2-3um latis, laevis, conidiophoris secundariis ex
hyphis secundariis lateraliter productis, solitariis.
Conidiis anguste cylindraceis vel filiformibus, pallidissime
brunneo-olivaceis, leniter curvatis, 3-6 septatis, ad apicem
mbotundatis;, basi leniter truncatis, 30=60 x 2.5-3m.
Habitat in foliis vivis Vigna sp. (Leguminosae), ad
Raiganj, Dinajpur occidentalis, Bengal occidentalis, India,
wed. Bb. Ks Das, 3 XIe 1960, No. Pec4451 (Herb. LAM, Yen
#10582), typus.
at
Leaf spot indistinct. Caespituli hypophyllous, invisible
even under the hand lens. Primary mycelium internal:
hyphae very pale olivaceous, smooth, septate, branched,
2-3.5ym wide. Stromata well developed, dark brown,
globular, 20-40um in diameter. Conidiophores very short,
numerous in a dense or very dense fascicle, simple,
flexuous, O-l septate, O-2 geniculate, apex irregularly
rounded, conidial scars not visible, 22-75 x 3-5um.
Secondary mycelium superficial: hyphae pale brown-
Olivaceous smooth, septate, branched, 2-3ym wide, producing
laterally the solitary, secondary conidiophores. Conidia
narrowly cylindric or filiform, very pale brown-olivaceous
slightly curved, 3-6 septate, with a rounded apex and a
truncate base, not constricted, 30-60 x 2.5-3um.
On Vigna sp. (Leguminosae), in Raiganj, West Dinajpur, West
Bengal, India. Leg. B. K. Das, 3 November 1980, No. Pcc4451
(Herb. Lam, Yen!#10582).
2. Phaeoisariopsis argyreiae Yen, Kar & Das, sp. nov.
CFaigi 2
Maculis primo indistinctis, deinde brunneis et visibilis,
orbicularis vel suborbicularis, margine indistinctis, usque
7mm diam., dispersis vel confluentibus. Caespitulis
districte hypophyllis, atro-hirsutis. Mycelium primarium
immersum: hyphis pallidissime olivaceis, septatis, ramosis,
2-4um latis. Stromatibus atro-brunneis, irregularis, 40-60.um
altis, 25-40um latis. Conidiophoris districte hypophyllis,
pallide brunneo-olivaceis, laevis, 3-25 fasciculatis,
synnematiforme aggregatis, multiseptatis, simplicibus,
inferne leniter undulatis, superne leniter erectis et
separabilis, ad apicem rotundatis, cicatricibus conidialis
indistinctis, 60-350 x 4-5.5um. Conidiis pallide olivaceis,
cylindraceis, semper valide curvatis, 1-7 septatis, non
constrictis, ad apicem rotundatis basi attenuatis et
truneatias, 55-110 x 6-7um.
Habitat in foliis vivis Argyreia roxburgnii Choisy
(Convolvulaceae), ad Raiganj, Dinajpur occidentalis, Bengal
Sccagentalis, india, Leg. BwaK. *Das, (12° xX 1980) No. Pcecd492
(Herb. LAM, Yen #10588), typus.
Leaf spot at first indistinct, only some rounded chlorosis
area, later becoming brown and visible, orbicular or
suborbicular, with an indistinct margin, up to 7mm in
diameter, scattered or sometimes confluent. Caespituli
strictly hypophyllous, visible under hand lens in dark hairy
form. |~ Primary mycelium internal: hyphae very pale
olivaceous, septate, branched, 2-4um wide. Stromata
well-developed, dark brown, irregular, 40-60um high and
Fig. 1. Cercoseptoria vignicola Yen, Kar & Das: Fascicles
of primary conidiophores and formation of external hyphae;
B, External hyphae and formation of secondary conidiophores;
Gr conidia.
38
oO”
25-40um wide. Conidiophores strictly hypophyllous, pale to
medium olivaceous, synnematous, 3-25 in loose or dense
fascicles, smooth, multiseptate, simple, slightly sinuous
below, substraight above, with a rounded apex, conidial
cicatrices indistinct, 60-350 x 4-5.5um. Conidia pale
oOlivaceous, cylindric, always curved, smooth, with a broadly
rounded apex, 1-7 septate, not constricted, base attenuate
and truncate, 55-110 x 6-7um.
On Argyreia roxburghii Choisy (Convolvulaceae), in Raiganj,
West Dinajpur, West Bengal, India. Leg. B. K. Das, 12
October 1980, No. Pcc4492 (Herb. LAM, Yen #10588).
3. Pseudocercospora alternantherae Yen, Kar & Das, sp. nov.
PAGS
Maculis amphiphyllis, ovoideis vel fusiformis,
griseo-brunneis, margine aliquantum indistinctis, saepe
dispersis vel confluentibus, 3-12mm longis, 2-4mm latis.
Caespitulis amphigenis, autem plerumque hypophyllis,
invisibilis. Mycelium primarium immersum: hyphis
olivaceo-brunneis, septatis, ramosis 2-5um latis.
Stromatibus brunneis vel atro-brunneis, globosis 20-40um
diam. Conidiophoris primariis amphiphyllis, autem in
hypophyllo plus abundis, 5-58 in fasciculo per stomatibus
emergentis, pallide burnneo-olivaceis, simplicibus, fere
flexuosis, laevis, 0-3 septatis, 0-2 geniculatis, ad apicem
rotundatis vel attenuatis, 15-55 x 3-4.5um. Mycelium
secundarium superficiale: hyphis ex stomatibus oriundis vel
cum conidiophoris primariis commixa in fasciculo per
stomatibus emergentis, pallide brunneo-olivaceis, repentis,
septatis, ramosis, 2-3.5um latis, conidiophoris secundariis
lateraliter gerentis. Conidiis obclaviformibus vel
obclavato-cylindraceis, pallide brunneo-olivaceis, rectis
vel leniter curvatis, laevis, 3-10 septatis, ad apicem
rotundatis, basi obconico-attenuatis et truncatis, 32-90 x
2.5-4ym.
Habitat in foliis vivis Alternanthera sp. (Amarathaceae), ad
Purulea, Bengal occidentalis, India, leg. B. K. Das, 7 I
1981, No. Pcc4483 (Herb. LAM, Yen #10583), typus.
Leaf spot amphigenous, ovoid or fusiform, brown grayish with
an indistinct margin, scattered, but often confluent, 3-12mm
long and 2-4mm wide. Caespituli amphiphyllous, but mostly
hypophyllous, invisible even under hand lens. Primary
mycelium internal: hyphae brown olivaceous, septate,
branched, 2-5um wide. Stromata brown or dark brown,
globular, 20-40um in diameter. Primary conidiophores
amphigenous, but more abundant on the lower surface, 5-58 in
fascicles emerging through the stomata, pale brown
Olivaceous, simple, generally flexuous, smooth, 0-3 septate,
0-2 geniculate, apex rounded or attenuate and shouldered,
Fig. 2. Phaeoisariopsis argyreiae Yen, Kar & Das: A,
Fascicles of old conidiophores; B, Stromata and young
conidiophores; C, Conidia.
40
a
a
Noy,
Ve:
, N
4l
15-55 x 3-4.5um. Secondary mycelium external: hyphae
arising from the stomata or from a stromata and mixed with
the primary conidiophores in a regular fascicle, pale brown
Olivaceous, repent, septate, branched, 2-3.5um wide, bearing
the secondary conidiophores as lateral branches which are
similar to the primary conidiophores. Conidia obclaviform
or obclavato-cylindric, pale brown olivaceous, straight or
slightly curved, smooth, 3-10 septate, apex rounded, base
obconic-attenuate and truncate, 32-90 x 2.5-4ym.
On Alternanthera sp. (Amaranthaceae), in Purulea, West
Bengal, India. Leg. B. K. Das, 7 January 1981, No Pcc4483
(Herb. LAM, Yen #10583).
Note: The Pseudocercospora alternanthericola (Pavgi & U. P.
Singh) Deighton differs from this fungus by its mostly
epiphyllous fruiting and thicker conidia (2.4-6.3um compared
with 2.5-4um). On the other hand, the Cercospora
alternantherae-nodiflorae Sawada differs from this species
by its leaf spot indistinct or none.
4. Pseudocercospora daturina (Yen) Deighton. CMI Mycol.
Pap. 140: 143, 1976.
=Cercospora daturina Yen. Rev. de Mycol. 30: 171, 1965.
Leaf spot indistinct, sometimes as chlorosis area without
definite margin. Caespituli amphiphyllous, grayish, effuse.
Primary mycelium internal: hyphae almost colorless, septate,
branched, 1.5-2.5um wide. Stromata absent. Primary
conidiophores amphiphyllous, 12-40 in fascicles, simple when
young, but generally branched when old, pale brown
Olivaceous, flexuous, 1-4 (rarely 5-6) septate, 1-3
geniculate, sometimes with constrictions at the septum, apex
attenuate and decorated with a conidial cicatrice (1.5ym in
diameter), 30-70 x 3.5-6ym (Yen: 30-80 x 4-6um). Secondary
mycelium superficial: hyphae arising from the stromata,
pale olivaceous, repent, septate, branched, 2-3.5um wide,
bearing the solitary secondary conidiophores as lateral
branches. Conidia obclavate-cylindric, pale brown
Olivaceous, straight or slightly curved, sometimes slightly
Sinuous, 3-10 septate, apex subconic, base
attenuate-truncate, 50-117 x 3.5-5Sum (Yen: 51-123 x
3.5-5um).
On Datura atramonium L. (Solanaceae), in Baluhati, Hocorah,
West Bengal, India. Leg. B. K.. Das, 22 August 1980, No.
Pcc4323 (Herb. LAM, Yen #10579).
Distribution: India and Singapore
Fig. 3. Pseudocercospora alternantherae Yen, Kar & Das: A,
Fascicle of conidiophores and formation of external hyphae;
B, Young conidiophores; C, Old conidiophores, D & E,
Formation of secondary conidia; F, Conidia. 5.
42
5. Pseudocercospora euphorbiae-piluliferae Yen, Kar & Das
Sp. nov. 19. , A-D )
Maculis hypophyllis, minusculis, obscure brunneis, margine
indistinctis, usque 2mm diam. in epiphyllo visibilis, in
hypophyllo indistinctis. Caespitulis hypophyllis, raro
amphiphyllis, non aspectabilis. Mycelium primarium
immersum: hyphis subhyalinis, septatis, ramosis, 2-3.5ym
latis. Stromatibus nullis. Conidiophoris plerumque
hypophyllis, pallide brunneo-olivaceis, 2-5 in fasciculo per
stomatibus emergentis, simplicibus vel ramosis, laevis,
erectis vel leniter flexuosis, aliquandum undulatis, 0-6
septatis, non geniculatis, cicatricibus conidialis
indistinctis, ad apicem rotundatis, 15-80 x 3.5-5um.
Conidiis obclaviformibus vel obclavato-cylindraceis, pallide
brunneo-olivaceis, rectis vel leniter curvatis, 3-5
septatis, non constrictis, ad apicem rotundatis, basi
truncatis, (338-745 x 3—5im,
Habitat in foliis vivis Euphorbiae piluliferae L.
(Euphorbiaceae), ad Mashlandapur, Nadia, Bengal
occidentalis; India|) Leq. Bay K. Das,/i3 Vis 19eo ie.
Pcc4305 (Herb. LAM, Yen #10584), typus.
Leaf spot small, brown with an indistinct margin, visible on
upper surface, but invisible on lower surface, up to 2mm in
diameter. Caespituli mostly hypophyllous, rarely
amphiphyllous, invisible. Primary mycelium internal: hyphae
almost colorless, septate, branched, 2-3.5um wide. Stromata
absent. Conidiophores generally hypophyllous, 2-5 in
fascicles emerging through a stomata, pale brown olivaceous,
simple or branched, smooth, with a slightly undulated
membrane, straight or slightly flexuous, O-6 septate, not
geniculate, conidial scars indistinct, apex rounded, 15-80 x
3.5-5um. Conidia obclavate or obclavate-cylindric, pale
brown olivaceous, straight or slightly curved, 3-5 septate,
not constricted, apex rounded, base truncate, 38-74 x 3-5um.
On Euphorbia pilulifera L. (Euphorbiaceae), in Mashlandapur,
Nadia, West Bengal, India. Leg. B. K Das, 13 August 1980,
No Pcc4305 (Herb. LAM, Yen #10584).
Note: This fungus differs from others (Cercospora petila
Thir. & Chupp, C. euphorbiae-pubescentis Unam. and C.
euphorbiaecola Tharp.) on Euphorbia, by its small brown leaf
spots, hypogenous caespituli, poor fascicles and absence of
stromata.
6. Pseudocercospora pantoleuca (Saccardo) Deighton CMI
Mycol. Pap. 11402507, 97Gemn Figse > )
=Cercospora pantoleuca Saccardo, nom. nov. (as 'Syd.
& Sacc.') in Sdccardo & Trotter, Syll. Fung. 25:906)
LOS 1s
=Cercospora pantoleuca H. Sydow & P. Sydow, Philipp. J.
Sci Sec iGUiBot “je e284-" 19137
NON Cercospora pantoleuca Saccardo, Michellia 1:268, 1879.
43
Leaf spot distinct, amphiphyllous, at first small (1-3mm in
diam.), vein-limited, green brownish, soon becoming grayish-
white in the center with a linear, slightly raised brown-
purplish margin, often 3-5 confluent in large blotches, up
to 5mm in diameter. Caespituli generally hypophyllous,
sometimes slightly amphigenous, dark punctiform. Mycelium
internal: hyphae very pale olivaceous, septate, branched,
2.5-4um wide. Stromata not well-developed, dark brown,
irregularly globular, 10-25ym in diameter. Conidiophores
chiefly hypogenous, 5-15 in fascicles emerging through the
stomata, pale brown olivaceous, simple, sometimes branched,
flexuous or sinuous, 1-6 septate, O-2 geniculate, not
continuous, sometimes slightly constricted, apex rounded or
attenuate and shouldered, conidial cicatrices indistinct,
25-55 x 3.5-4um. Conidia obclavate-cylindric, straight or
slightly curved, 3-11 septate, pale olivaceous, apex
rounded, base obconic-truncate.
On Clitoria ternatea (Leguminosae), in Palpara, Nadia, West
Benga sindia. \ Leg. B. K.. Das, 25 August 1980, No Pec4327
(Herb. LAM, Yen #10581).
Distribution: Philippines and India.
Note: The material of this species from West Bengal (India)
differs from others from the Philippines by the largely
hypophyllous fruiting and the conidiophores not being
continuous.
7. Pseudocercospora polyalthiae Yen, Kar & Das, sp. nov.
Maculis indistinctis. Caespitulis hypophyllis, invisibilis.
Mycelium primarium immersum: hyphis subhyalinis, septatis,
ramosis, 1.5-3um latis. Stromatibus nullis. Conidiophoris
solitariis, non fasciculatis, pallide brunneo-olivaceis,
sursum pallidioribus, plerumque simplicibus, interdum
ramosis, 0-8 septatis, O-3 geniculatis, apex attenuatis et
truncatis, interdum irregulariter rotundatis, cicatricibus
conidialis invisibilis, 12-70 x 3.5-5Sym. Mycelium
secundarium superficiale: hyphis repentis, pallide
Olivaceis, laevis, septatis ramosis, 1,5-3um latis,
conidiophoris lateraliter manifestibus. Conidiis gracilis,
obclavato-cylindraceis, pallide brunneo-olivaceis, rectis
vei leniter curvatis; 3-13 septatis, non constrictis,) ad
apicem rotundatis vel subconicis, basi obconico-truncatis,
40-£56) 26 2..5=-3 55 um.
Habitat in foliis vivis Polyalthiae suberosa Benth. &
Hooker (Annonaceae), ad Garia, 24-Parganas, Bengal
cceideritalis, India. Leg. 9B. K. Das, 15 XII 1979; No.
Pcc4095 (Herb. LAM, Yen #10585), typus.
Leaf spot indistinct or none. Caespituli hypophyllous,
invisible. Primary mycelium internal: hyphae very pale
45
olivaceous, septate, branched, 1.5-3um wide. Stromata
absent. Conidiophores solitary, not in fascicles, generally
simple, sometimes branched, straight or flexuous, pale brown
Olivaceous, paler towards the tip, 0-8 septate, 0-3
geniculate, apex attenuate and shouldered, conidial scars
HOWMaTStinck,. 12-70 x 3s5—5um. © Conidia narrowly
oblclavate-cylindric, pale brown olivaceous, straight or
slightly curved, 3-13 septate, not constricted, apex rounded
or subconical, base obconic-truncate, 40-156 x 2.5-3.5um.
On Polyalthia suberosa Benth. & Hooker (Annonacae), in
Garia, 24-Parganas, West Bengal, India. Leg. B. K. Das, 15
December 1979, No Pcc4095 (Herb. LAM, Yen #10585).
8. Pseudocercospora timorensis (Cooke) Deighton CMI Mycol.
Pap. 140:154, 1976.
=Cercospora timorensis Cooke Grevillea 12:38, 1883.
On Ipomoea batatas Lamx. (Convolvulaceae), in Samali,
24-Parganas, West Bengal, India. Leg. B. K. Das, 2
September 1980, No Pcc4345 (Herb. LAM, Yen #10580).
Distribution: Africa, China, Ecuador, India, Japan, Panama,
Philippines, and Singapore.
9. Pseudocercospora trematis-orientalis (Sun) Deighton CMI
PycolmPapil40s 155, Lovo." lye
=Cercospora trematis-orientalis Sun J. Agric. (Formosa)
PAiwane Prov. Coll. .9348 (reprint), 1955.
beat spot distinct, at first small, brown, “irregularly
circular, isolate, 0.5-lmm in diameter, soon extending into
large lesions, irregularly angular, vein-limited, gray-white
in the center, having a brown-purple border with indefinite
margin, more distinct on lower surface, scattered, up to 3mm
in diameter. Caespituli amphiphyllous, but chiefly
epiphyllous, dark punctiform on the upper surface. Mycelium
internal: hyphae very pale olivaceous, septate, branched,
2-4um wide. Stromata dark brown, globular, 20-40um in
diameter. Conidiophores 5-28 in fascicles emerging through
the stomata, pale brown-olivaceous, paler towards the tip,
straight or slightly flexuous, simple, not branched, smooth,
1-3 septate, 0-2 geniculate, apex rounded or attenuate,
conidial scars sometimes distinct, 40-95 x 3.5-4um. Conidia
pale brown-olivaceous, obclavate-cylindric, often slightly
curved, or slightly undulated, 3-8 septate, apex rounded or
subconical, base obconic-truncate, 40-108 x 3.5-4um.
On Trema orientalis Bl. (Urticaceae), in Raiganj, West
Dinajpur, West Bengal, India. leg. B. K. Das, 20 October
1980, No. Pcc4447 (Herb. Lam, Yen #10575).
Fig. 4. A-D, Pseudocercospora euphorbiae-piluliferae
Yen, Kar, & Das: A & D, Fascicles Of conidiophores; B, Young
conidiophores; C, Conidia. E-G, P. polyalthiae Yen,
Kar & Das: E, External hyphae and formation of secondary
conidiophores; F, Branched conidiophores; G, Conidia.
46
47
Distribution: Taiwan (China) and West Bengal (India).
Note: Sun proposed a common name, "white stars disease on
leaves of Trema", which is apropos because of the
symptomatic isolated white angular leaf spots which
characterize the infected leaves.
10. Pseudocercospora vignigena Yen, Kar & Das, sp. nov.
Eid. oO
Maculis distinctis, orbicularis vel irregulariter
orbicularis, clare brunneis, margine atro-brunneis,
dispersis, non confluentibus, 2-5mm diam. Caespitulis
amphiphyllis, nigrostrigosis. Mycelium primarium immersum:
hyphis fere incoloris, septatis, ramosis, 2-3.5um latis.
Stromatibus mediocoriter evolutis, atro-brunneis, globosis
vel irregulariter globosis, 15-40ym diam. Conidiophoris
primariis usque 60 in fasciculo per stomatibus emergentis,
pallide brunneo-olivaceis, laevis, erectis vel leniter
flexuosis, 1-3 septatis, 0-2 geniculatis, apex attenuatis et
Cubpigeeyercicatricibus conidialis distinctis, 22-75 \x 3-Sum.
Mycelium secundarium superficiale: hyphis ex cellulis
basalibus conidiophorum primariorum aliquorum oriundis,
repentis, pallide olivaceis, septatis, ramosis, 2.5-4um
latis, conidiophoris secundariis erectis lateraliter
gerentis. Conidiis cylindraceis, interdum
obclavato-cylindraceis, pallide olivaceis, rectis vel
leniter curvatis, 3-6 septatis, apice rotundatis, basi
attenuatis et truncatis, 33-60 x 4-5.5(-6)um.
Habitat in foliis vivis Vigna sp. (Leguminosae), ad Raiganj,
Dinajpur occidentalis, Bengal occidentalis, India. Leg. B.
K. Das, 3 XI 1980, No. Pcc4451 (Herb. LAM, Yen #10582),
typus.
Leaf spot distinct, orbicular or irregularly orbicular,
bright brown with a dark brown margin, scattered, more
visible on the upper surface, 2-5mm in diameter. Caespituli
amphiphyllous, visible under hand lens as short black hairs.
Primary mycelium internal: hyphae almost colorless,
septate, branched 2-3.5um wide. Stromata not well
developed, dark brown, globular or irregularly globular,
15-40um in diameter. Primary conidiophores up to 60 in
dense fascicles emerging through a stromata, pale
brown-olivaceous, smooth, simple, straight or slightly
flexuous, 1-3 septate, 0-2 geniculate, apex attenuate or
shouldered, conidial scars-distinct, 22-75 x 3-5um.
Secondary mycelium superficial: hyphae arising from the
base of some of the primary conidiophores, repent or lax
arcuate, pale olivaceous, septate, branched, 2.5-4um wide,
bearing secondary conidiophores as erect lateral branches.
Conidia generally cylindric, sometimes obclavate-cylindric,
pale olivaceous, straight or slightly curved, 3-6 septate
Fig. 5. Pseudocercospora pantoleuca (Saccaardo) Deighton:
A, Fascicles of young conidiophores; B & C, Fascicles of old
conidiophores; D, Branched conidiophores; F, Conidia.
48
49
apex rounded, base attenuate-truncate, 33-60 x 4-5.5(-6)um.
On Vigna sp. (Leguminosae), in Raiganj, West Dinajpur, West
Bengal, India. Leg. B. K. Das, 3 November 1980, No Pcc4451
(Herb. LAM, Yen #10582).
Note: This fungus differs from others (Pseudocercospora
cruenta (Saccardo) Deighton, Pe dolichs Evins
Everhart) Yen, BP mungo Deighton, and P.
Vignee-reticulatae Deighton fon Vigna by its distinct
leaf spots with definite margin and very dense fascicles
with up to 60 conidiophores.
ll. Pseudocercospora vitigena Yen, Kar & Das, sp. nov.
G oS ire iy gs RoE}
Maculis indistinctis. Caespitulis amphiphyllis, autem
principaliter hypophyllis, invisibilis. Mycelium primarium
immersum: hyphis subhyalinis, septatis ramosis, 1.5-2.5 um
latis. Stromatibus atro-brunneis, globosis vel
irregulariter globosis, 10-30um diam. Conidiophoris
primariis numerosis vel paucis in fasciculo per stomatibus
emergentis, pallide olivaceis, eretis vel leniter flexuosis,
simplicibus vel ramosis, laevis, continuis vel 1-2 septatis,
non geniculatis, apice rotundatis vel irregulariter
rotundatis, 10-30 x 2-3um. Mycelium secundarium
superficiale: hyphis ex stromatibus oriundis, pallidissime
olivaceis, repentis vel arcuatis, laevis, septatis,
conidiophoris secundariis lateraliter gerentis. Conidiis
pallide olivaceis, obclavato-cylindraceis, rectis vel
leniter curvatis, laevis, non constrictis, 3-11 septatis, ad
apicem rotundatis vel subrotundatis, basi
obconico-truncatis, 36-82 x 2.5-3.5um.
Habitat in foliis vivis Vitis sp. (Vitidaceae), ad Sylva
Hemtabad, Dinajpur occidentalis, Bengal occidentalis, India.
Leg. B./K. Das, 10 X 1980, No. Pcc4418 (Herb. LAM, Yen
#10577), typus.
Leaf spot indistinct or none. Caespituli amphigenous, but
chiefly hypophyllous, not visible even under hand lens.
Primary mycelium internal: hyphae very pale olivaceous,
septate, branched, 1.5-2.5um wide. Stromata slightly
developed, dark brown, globular or irregularly globular,
10-30um in diameter. Primary conidiophores emerging through
the stomata, very numerous in each of the larger fascicles,
but few in smaller fascicles, pale olivaceous, straight or
slightly flexuous, simple or branched, smooth, continuous or
1-2 septate, not geniculate, apex rounded or irregularly
rounded, 10-30 x 2-3ym. Secondary mycelium external:
hyphae arising from the stomata, sometimes from stromata,
Fig. 6. Pseudocercospora vignigena Yen, Kar & Das: A,
Young conidiophores; B, Fascicles of old conidiophores; C,
External hyphae and formation of secondary conidiophores; E,
Conidia.
51
repent or arcuate, very pale olivaceous, smooth, septate,
bearing secondary conidiophores as lateral branches.
Conidia very pale olivaceous, obclavate-cylindric, straight
Or slightly curved, smooth, not constricted, 3-11 ‘septate,
apex rounded or subrounded, base obconic-truncate, 36-82 x
2.5-3.5um.
On Vitis sp. (Vitidaceae), in Hemtabad forest, West
Dinajpur, West Bengal, India. Leg. B. K. Das, 10 October
1980, No Pcc4418 (Herb. LAM, Yen #10677).
Note: The indistinct leaf spots, amphigenous fruiting, and
narrow pale olivaceous conidia separate this species from
ouners oni Vitis.
12. Stenetla canthii Yen, Kar & Das, sp. nov. (Fig. 7, F&G)
Maculis distinctis, orbicularis vel irregulariter
orbicularis, brunneo-albis et zona lata obscure brunnea
restrictis, 2-10 mm diam., interdum confluentibus, usque
17mm longis. Caespitulis districte hypophyllis, invisibilis.
Mycelium primarium immersum: hyphis pallidissime olivaceis,
septatis, ramosis, 1.5-3um latis. Stromatibus nullis.
Conidiophoris solitariis, ex hyphis secundariis oriundis,
brunneo-olivaceis vel pallide olivaceis, erectis vel leniter
flexuosis, simplicibus, laevis, 1-3 septatis, O-l
geniculatis, antice attenuatis, denticulatis et cicatricibus
atro-brunneis ornatis, 21-45 x 3-3.5ym. Mycelium
secundarium superficiale: hyphis ex stromatibus oriundis,
pallide olivaceis, repentis vel arcuatis, subtiliter
verruculosis, septatis, conidiophoris solitariis lateraliter
gerentis. Conidiis plerumque cylindraceis, interdum
obclavato-cylindraceis, pallide olivaceis, subtiliter
verruculosis, plerumque curvatis, raro rectis, 1-7 septatis,
ad apicem rotundatis, inferne attenuatis et in hilum
truncatum, 18-112 x 3-4.5um.
Habitat in foliis vivis Canthii dedymi Roxb. (Rubiaceae), ad
Baraipur, 24-Parganas, Bengal occidentalis, India. Leg. B.
K. Das, 15 IX 1980, No. Pcc4374 (Herb. LAM, Yen #10590),
typus.
Leaf spot distinct, brown whitish, with a wide indistinct
dark brown margin, 2-10mm in diameter, sometimes confluent
up to 17mm long. Caespituli strictly hypophyllous,
invisible. Primary mycelium internal: hyphae very pale
olivaceous, septate, branched, 1.5-3um wide. Stromata
absent. Conidiophores arising from secondary hyphae as
lateral branches, brown olivaceous or pale brown olivaceous,
straight or slightly flexuous, simple, smooth, 1-3 septate,
Fig. 7. A-E, Pseudocercospora vitigena Yen, Kar & Das:
A, Young conidiophores; B, Branched conidiophores; C, Poor
fascicles of conidiophores; D, External hyphae and formation
of secondary conidiophores; E, Conidia. F-G, Steneila
canthii Yen, Kar & Das: F, External hyphae and formation
of normal conidiophores; G, Conidia.
52
Do
not geniculate, apex attenuate, denticulate and decorated
with dark brown conidial ciccatrices, 21-45 x 3-3.5ym.
Secondary mycelium external: hyphae arising from stomata,
pale olivaceous, repent or arcuate, finely verruculose,
septate, bearing laterally the normal conidiophores.
Conidia generally cylindric, sometimes obclavate-cylindric,
pale olivaceous, finely verruculose, slightly curved,
sometimes straight, 1-7 septate, apex rounded, base
attenuate-truncate, 18-112 x 3-4.5um.
On Canthium dedymum Roxb. (Rubiaceae), in Baraipur,
24-Parganas, West Bengal, India. Leg. B. K. Das, 15
September 1980, No Pcc4374 (Herb. LAM, Yen #10590).
13. Stenella dioscoreicola Yen, Kar & Das, sp. nov. (Fig. 8)
Maculis irregulariter orbicularis, brunneis, margine
indistinctis, dispersis, 3-12mm diam. Caespitulis districte
hypophyllis, in hypophyllo obscure punctiformibus. Mycelium
primarium immersum: hyphis pallidissime olivaceis,
septatis, romosis, 1.5-3.5um latis. Stromatibus
atro-brunneis, irregulariter globosis, 24-40um diam.
Conidiophoris hypophyllis, 12-40 in fasciculo per stomatibus
emergentis, pallide brunneo-olivaceis, sursum pallidioribus,
Simplicibus vel raro ramosis, cylindraceis, erectis vel
leniter flexuosis, O-3 septatis, non geniculatis, ad apicem
PoLundatis, CiGapricipus conidialis:indistinctis,. 4/—-60 x
4-5yum. Mycelium secundarium superficiale: hyphis parvum
explicatis, cum conidiophoris primariis commixa in fasciculo
per stomatibus emergentis, pallide olivaceis, repentis,
septatis, ramosis, subtiliter verruculosis, 2-3,m latis,
conidiophoris secundariis lateraliter raro gerentis.
Conidiis pallide brunneo-olivaceis, obclaviformis vel
obclavato-cylindraceis, plerumque curvatis vel undulatis,
subtiliter verruculosis, 3-9 septatis, apice obtusis vel
rotundatis, basi obconico-truncatis, 50-130 x 3.5-4.5(-5) um.
Habitat in foliis vivis Dioscoreae sp. (Dioscoreaceae), ad
Sylva Chelapata Cooch Behar, Bengal occidentalis, India.
beg. B.\K. Das; 17 X-1980, Pcc4452 (Herb. LAM, Yen #10578),
typus.
Leaf spot irregular orbicular, brown with an indistinct
margin, scattered, 3-12mm in diameter. Caespituli strictly
hypophyllous, visible under hand lens as dark punctiform.
Primary mycelium internal: hyphae very pale olivaceous,
septate, branched, 1.5-3.5um wide. Stromata dark brown,
irregularly globular, 24-40um in diameter. Conidiophores
hypophyllous, 12-40 in fascicles, arising from a stomata,
generally simple, rarely branched, cylindric, straight or
slightly flexuous, pale brown olivaceous, paler towards the
tip, 0-3 septate, not geniculate, apex rounded, conidial
Fig. 8. Stenella dioscoreicola Yen, Kar & Das: A,
Fascicles of primary conidiophores; B, Branched conidio-
phores; C, Young conidiophores; D, External hyphae and
formation of secondary conidiophores; E, Conidia.
54
39
scars indistinct, 17-80 x 4-5um. Secondary mycelium
superficial: hyphae very poorly developed, mixing with the
primary conidiophores and arising from a stomata, pale
olivaceous, repent, septate, branched, finely verruculose,
2-3um wide, bearing occasionally the secondary conidiophores
as lateral branches. Conidia pale brown olivaceous,
obclaviform or obclavate-cylindric, generally curved or
undulated, finely verruculose, 3-9 septate, apex obtuse or
rounded, base obconici-truncate, 50-130 x 3.5-4.5(-5) um.
On Dioscorea sp. (Dioscoreaceae), in Forest of Chelapata,
Cooch Behar, West Bengal, India. Leg. B. K. Das, 17 October
1980, No Pcc4452 (Herb. LAM, Yen #10578).
14. Stenella myxa (Sydow in Sydow & Mitter) Yen, Kar & Das
combs*)nov.!\ (Fig. 9, A=C)
=Cercospora myxa Sydow in Sydow and Mitter, Ann. Mycol.
33200; L935.
Leer spot indistinctyor none... Caespitul lL stricely
hypophyllous, in effuse dark olivaceous irregular patches.
Primary mycelium internal: hyphae almost colorless,
septate, branched, 1.5-3um wide. Stromata not developed.
Conidiophores hypophyllous, solitary, arising as lateral
branches on the external mycelial hphae, pale brown
Olivaceous, simple continuous or 1-2 septate (rarely 3),
smooth, substraight below, sinuous or subgeniculate towards
the tip, apex attenuate and rounded, 13-26 x 5-6um. Conidia
cylindric, brown olivaceous or pale brown olivaceous,
straight or slightly curved, finely verruculose, 2-14
septate (Chupp:2-12 septate), apex rounded, base attenuate
and truncate, 24-110 x 5-6um.
On Cordia myxa L. (Boraginaceae), in Kamarkundu, Nnoia, West
Bengal, India. Leg. B. K. Das, 23 February 1980, No Pcc4234
(Herb. LAM, Yen #10587).
Note: We have sent the material of this species to the
Commonwealth Mycological Institute in Kew, England, and it
has been identified as a Stenella sp. According to the
description of Cercospora myxa Sydow in Sydow and Mitter,
which parasitizes also Cordia myxa in India, we find it is
similar to our fungus.
15. Stenella stephaniae Yen, Kar & Das, sp. nov. (Fig. 9,
D-G)
Maculis brunneis, angularis, per nervuli limitatis, in
epiphyllo plus visibilis, dispersis, 1-2mm diam.
Caespitulis hypophyllis invisibilis. Mycelium primarium
immersum: hyphis fere incoloris, septatis, ramosis,
2.5-4.5um latis. Stromatibus atro-brunneis, globosis
Fig. 9. A-C, Stenella myxa (Sydow in Sydow & Mitter)
Yen, Kar & Das: A, External hyphae and formation of
conidiophores; C, Conidia. D-G, Stenella stephaniae Yen,
Kar & Das: D, Young conidiophores; E, Conidiophores and
formation of conidia; F, Branched conidiophores; G, Conidia.
56
10-45um diam. Conidiophoris hypophyllis, ex stomatibus
Ooriundis, 2-8 in fasciculotis, pallide brunneo-olivaceis,
Ssimplicibus, interdum ramosis, laevis, erectis, 1-4
septatis, apex rotundatis vel attenuatis et denticulatis,
cicatribus sporarum atro-brunneis manifestibus, 25-120 x
4-5um. Conidiis breviter cylindraceis, pallide
brunneo-olivaceis, leniter curvatis, 1-3 septatis, ad apicem
rotundatis, basi attenuatis et truncatis, 31-45 x 5-7um.
Habitat in foliis vivis Stephania hernandifolia Walp.
(Menispermaceae), ad Ragan}, Dinajpur occidentalis, Bengal
occidentalis, India. Leg. B. K: Das, 248xX% 1979, No.3 Frce47an
(Herb. LAM, Yen #10591), typus.
Leaf spot small, brown, angular and vein-limited, more
distinct on upper surface, scattered, 1-2mm in diameter.
Caespituli strictly hypophyllous, invisible even under hand
lens. Primary mycelium internal: hyphae almost colourless,
septate, branched, 2.5-4.5um wide. Stromata dark brown,
globular, 10-45um in diameter. Conidiophores hypophyllous,
emerging through the stomata, 2-8 in poor fascicles, rather
pale olivaceous and paler towards the tip, smooth, usually
simple, occasionally branched, straight below, slightly
flexuous above, 1-4 setate, not geniculate, apex rounded or
attenuate and denticulate, dark-brown cicatrices conidial
decorated on the shoulders, 25-120 x 4-5um. Conidia shortly
cylindric, pale brown-olivaceous, slightly curved, 1-3
septate, apex rounded, base attenuate- truncate, 31-45 x
5-7um.
On Stephania hernandifoloa Walp. (Meninspermaceae), in
Raganj, Wet Dinajpur, West Bengal, India. Leg. B. K. Das,
24 October 1979, No Pcc 4231 (Herb. LAM, Yen #10591).
Note: We have sent the material of this fungus to the
Commonwealth Mycological Institute in Kew, England, and it
has been identified as a Stenella sp. Although CMI has not
mentioned the species name, we consider it as a new one.
LITERATURE CITED
Chupp, C. 1953. A monograph of ‘the genus Cercospora.
Ithaca, New York. 667 pp.
Deighton, F. C. 1976. Studies on Cercospora and allied
genera. 6. CMI Mycol. Pap. 140:1-168.
Deighton, F. C. 1979. Studies on Cercospora and allied
genera.’ 7. CMI) Mycol. Pap. 14471-—56.
Kar, A. K. and M. Mandal. 1970. New Cercospora spp. from
West’ ‘Bengal Iis7Trans. Brit. Mycol.’ Soce 542423-4338
Pavgi, M./S. "and U%" P.’-Singh. 1965:° Parasitic fungi strom
North: India. 5. Mycopath. Mycol. Appl. 27:89-96.
Sawada, K. 1928. Descriptive Catalog of Formosan Fungi. 4.
Rept... Dept. Agri. (Gov. Res.’ Inst. Formosa. 33. 123 Pp.
Sawada, K. 1959. Descriptive Catalog of Formosan Fungl.
ii). Cob. JAgri. Nat.Taiwan Univ. Spec. Publ 7S. (26130 p.
Sun, S. H. 1955. Studies on the genus Cercospora found
in Taiwan. I. J. Agric. (Formosa) Taiwan 4:38-48.
57,
SVdow,) .—..and) J. H. Mitter.. Fungi .indici. Il. Ann., Mycol.
33:46-71.
Yen, J. M. 1965. Etude sur les champignons parasites du
Sud-Est asiatigque III. Deuxieme note sur quelques
nouvelles especes de Cercospora de Singapour. Rev. de
Mycol. 30:166-204.
ACKNOWLEDGEMENT
We are grateful to Dr. F. C. Deighton and the Commonwealth
Mycological Institute, Kew, England, for help given in the
determination of critical specimens. We are much indebted
Lowpr. Don RR. Reynolds and Dr. M. M. > Thaung ‘of the Natural
History Museum, Los Angeles, California, for reviewing the
manuscript.
MYCOTAXON
WO OCA VE SENO. Ble ODD. (OB Hi9 October-December 1982
STUDIES ON HYPHOMYCETES FROM WEST BENGAL, INDIA, II.
CERCOSPORA AND ALLIED GENERA OF WEST BENGAL, 2
JO-MIN YEN
Natural History Museum, 900 Exposition Boulevard
Los Angeles, California 90007, U.S.A.
and
AéoK s(\KAR ‘and B. “kK. DAS
Department of Botany, Presidency College, Calcutta-700073, India
ABSTRACT
This study includes descriptions and illustrations of
nine new species and two new combinations of Indian
Cercosporae: Pseudocercospora brideliicola sp. nov., on
Bridelia sp.; P. micheliicola sp. nov., on Michelia
champaca; P. oroxyligena sp. nov., on Oroxylum indicum P.
pavettae-indicae (Gov. & Thirum.) comb. nov., on Pavetta
indica; P. stillingiae (Ell. & Ev.) comb. nov., on Sapium
sebiferum; P. tectonicola sp. nov., on Tectona grandis;
P. viticigena sp. nov., on Vitex negundo; Stenella coffeae
sp. nov., on Coffea bengalensis; S. garugae Sp. nov., on
Gargua pinnata; S. oroxylicola sp. nov., on Oroxylum
indicum; and S. xeromphigena sp. nov., on Xeromphis
uliginosa.
1. Pseudocercospora bridelticola. Yen, Kar G Das, mere
TOW ei( Ealsgiar |
Maculis indistinctis, Caespitulis hypophyllis, effusis, velutinis,
griseis, mox interdum confluentis et in inferiore superficie saepe
fere totam folii paginam omino obtegentis. Mycelium primarium
immersum: hyphis pallidissime olivaceis, septatis, ramosis, 3-4 um
latis. Stromatibus nullis vel parvis, substomatalis, ex hyphis paucis
et subhyalinis compositis. Conidiophoris 2-15 in fasciculo per
stomatibus emergentis, divergentis, laevis, olivaceis et sursum
pallidioris, simplicibus, interdum ramosis, erectis vel leniter
flexuosis, 2-10 septatis, 0-2 geniculatis, superne interdum denticu-
latis, ad apicem rotundatis vel attenuatis, cicatricibus conidialis
interdum visibilis (1.5-2.5 wm diam.), 40-180 (-250) x 5-6 um.
Mycelium secundarium superficiale: hyphis pallide olivaceis, ex
stomatibus et conidiophoris primariis oriundis, septatis, ramosis,
laevis, 2.5-3 um latis, conidiophoris secundariis lateraliter mani-
festibus. Conidiis obclavato-cylindraceis, pallide olivaceis, rectis
vel leniter curvatis, plerumque 3 septatis (raro 1 vel 4 septatis),
Sy
Fig. 1. Pseudocercospora brideliicola: A, Young conidiophores;
B, Rudimentary stromata; C, Formation of conidia; D, Formation of
secondary conidiophores; E, Conidia.
60
apice rotundatis, basi obconico-truncatis, 21-67 x 4.5-5 um.
Habitat in foliis vivis Brideliae sp. (Euphorbiaceae), ad Bibirhat,
24-Parganas, Bengal occidentalis, India, leg. B.K. Das, 10 I 1980, No.
PCC 4125 (IMI 250379) (Herb. LAM Yen #10601, holotypus).
Leaf spot indistinct or none, Caespituli hypogenous,
effuse, velutinous, grayish, sometimes extending and cover-
ing all the under surface of the leaf. Primary mycelium
internal: hyphae very pale olivaceous, septate, branched,
3-4 um wide. Stromata absent or very rudimentary, only a
loose mass of subhyaline internal hyphae under the stomata.
Conidiophores 2-12 in fascicles emerging through the sto-:
mata, olivaceous below and paler or subhyaline towards the
tip, simple, sometimes branched, 2-10 septate, 0-2 genicu-
late, sometimes denticulate above, apex rounded or attenu-
ate, conidial scars visible on the shoulder or at the end
of denticles, 40-180 (-250) x 5-6 wm. Secondary mycelium
superficial: hyphae pale olivaceous arising from the
stomata or from the prolongation of primary conidiophores,
septate, branched, 2.5-3 wm wide, often bearing the short
secondary conidiophores as lateral branches. Conidia
obclavate-cylindric, pale olivaceous, straight or very
slightly curved, generally 3 septate, rarely 1 or 4 septate,
apex rounded, base obconic and attenuate-truncate, 21-67 x
4.5-5 um.
On Bridelta sp. (Euphorbiaceae), in Bibirhat, 24-Parganas,
West, Bengal India, weg. BUKt Das. 10. 1 Los0s) Nov hCG aes
(IMI 250379)" (Herb. LAM Yen #10601):
Note: Sawada (1943) has described in Taiwan, Cercospora
atridis Syd., which parasitizes the leaves of Bridelta monotca
(Lour.) Merr.; it differs from this fungus by its dark-brown
wider conidia (7.5-8 wm compared with 4.5-5 wm). On the
other hand, Pseudocercospora brideliae Deighton differs from
this species by its much longer dark olivaceous conidiophores
(up to 450 um long) and by its condial septatum (3-15 septate
compared with 3 septate).
2. PSEUDOCERCOSPORA HIBISCI-CANNABINI (Sawada) Deighton
(Fig. 2) Mycol. Pap. 140:145, 1976 = cercospora hibisci-
cannabini Sawada, Descr. Cat. Formosan Fungi, 2:153, 1921.
Leaf spot indistinct, but some angular areas darkish,
vein-limited, more visible on lower surface, scattered, 0.5-
3 mm in diameter, sometimes confluent. Caespituli amphi-
genous, but chiefly hypophyllous, effuse, dark-gray, some-
times extensively velutinous on lower surface of the leaf.
Primary mycelium internal:hyphae subhyaline, septate, bran-
ched, 2.3-5 um wide. Stromata absent or very rudimentary.
Conidiophores emerging through the stomata, pale olivaceous,
forming dense to very dense fascicles: above the stomata,
pale olivaceous, forming dense to very dense fascicles above
the stomata, simple or branched, flexuous, warehy «strane,
1-7 septate, apex rounded, or attenuate-truncate and some-
times shouldered, 14-40 x 3-3.5 wm. Secondary mycelium
superficial: hyphae pale olivaceous, septate, branched, 2-4
um wide, bearing secondary conidiophores as lateral branches.
61
Conidia obclavate-cylindric, very pale olivaceous, straight
or slightly curved, 1-9 septate (Sawada: 3-9 septate), apex
reunded sepase obconic-truncate orratlenucate-truncate, 21-75
x 3=3.5 (-4) um (Sawada: 40-88 x 3-325 ym).
On Hibiscus rosa-sinensis L., (Malvaceae), in Raiganj,
Westubinaypur, West, Benvals india, deg. Bon, Das, 15) 1X 1980,
NoweeGC 3802 (IME -250373). (Herb. ;LAM Yen-#10599) .
Note: Pseudocercospora abelmoschi (Ell. §& Ev.) Deighton
differs from.this species, by its wider conidia (3-7 um
compared with 3-355.m).. But ald thevsystematic characters
Of this fungus are similar to those of Pseudocercospora
hibisci-cannabini (Sawada) Deighton, which have been well
described and illustrated by Sawada (1921).
Fig. 2. Pseudocercospora hibisci-cannabini: A, Old and branched
conidiophores; B, Formation of secondary mycelial hyphae; C & D,
Secondary conidiophores; E, Conidia.
62
3. Pseudocercospora micheliicola Yen, Kar §& Das, sp.
PO Vieed Odo 3)
Maculis distinctis, angularis, nervuli limitatis, interdum
irregularis, mediocriter brunneis, autem albo-griseis in medio, dis-
persis, 2-5 mn diam., in-inferiore superficie minus distinctis.
Caespitulis amphiphyllis, autem principaliter epiphyllis, punctiformis,
atro-brunneis, in superiore superficie distributis. Mycelium immersum:
hyphis pallide olivaceis, laevis, septatis, ramosis, 2.5-4.5 wm latis.
Stromatibus atrobrunneis, globosis vel subglobosis, 20-50 um diam.
Conidiophoris numerosis in dense vel valde dense fasciculo per
stomatibus emergentis, mediocriter olivaceis vel pallide olivaceis,
sursum pallidioris, laevis, simplicibus, semper tortuosis vel sinuosis,
0-3 septatis, 0-2 geniculatis, ad apicem rotundatis vel attenuatis,
Cicatricibus conidialis indistinctis, 10-55 x 4-5 um. Conidiis
obclaviformibus, olivaceis vel mediocriter olivaceis, rectis vel
curvatis, 4-10 septatis, laevis, non constrictis, apice subacutis vel
obtusis, basi obconico-truncatis, 55-120 x 5.5-7 um.
Habitat in foliis vivis Mtcheltae champacae L. (Magnoliaceae), ad
Rajahatkhawa Sylva, Alipurduar, Jalpaiguri, Bengal occidentalis, India,
leg. B.K. Das, 17 x 1980, No. PCC 4420 (Herb. LAM Yen #10603, holo-
typus) .
Leaf spot distinct, angular, vein-limited, sometimes
irregular, mid brown with a yrayish white center,, 2-5 mm in
diameter, scattered, more visible on upper surface.
Caespituli amphigenous, but chiefly epiphyllous, punctiforn,
black brown, loosely distributed over the spot on upper
surface. Mycelium internal: hyphae pale olivaceous, smooth,
septate, branched, 2.5-4.5 wm wide. Stromata well developed,
dark brown, globular, or subglobular, 20-50 um in diameter.
Conidiophores emerging through the stomata, numerous in
dense or very dense fascicles, mid olivaceous to pale
olivaceous, but very pale olivaceous or subhyaline towards
the tip, smooth, simple, always tortuous with a sinuous
membrane, 0-3 septate, 0-2 geniculate, apex rounded or
shouldered and attenuate, conidial scars not distinct, 10-
55 x 4-5 pm. Conidia obclaviform, olivaceous or mid
olivaceous, straight or curved, smooth, not constricted,
4-10 septate, apex subacute or obtsue, base obconic-truncate,
Son 20 Oe awe a
On Michelta champaca L. (Magnoliaceae), in Rajabhatkhawa
Forest, Alipurduar, Jalpaiguri, West Benyal. India, ler:
Bek. Das, 17 0X T9805 No. PCC. 4420 .(Herb.. PAN Venms1 060 4)e
Note: Cercospora micheliae Boedijn differs from this
fungus by its conidiophores only arising from the external
mycelial hyphae, without stromata, and especially by its
much narrower conidia (3.5-4.5 um wide compared with 5.5-7
um wide). q
63
Fig.3. Pseudocercospora micheliicola: A, Conidiophores and formation
of conidia; B, Young conidiophores; Cc, Old conidiophores and stromata;
D, Conidia.
64
4. Pseudocercospora oroxyligena Yen, Kar & Das, sp.nov.
(Figs 24)
Maculis indistinctis, tandem areae parvae angularis, nervuli limi-
tatis, rufo-brunneis, 2-5 mm diam., in inferiore superficie invisibilis.
Caespitulis districte hypophyllis, invisibilis. Mycelium immersum:
hyphis pallide olivaceis, laevis, septatis, ramosis, 2.5-4.5 um latis.
Stromatibus nullis. Conidiophoris solitariis vel 2-3 in fasciculo per
stomatibus emergentis, pallide olivaceis, sursum pallidioris (fere
hyalinis), simplicibus, erectis vel leniter curvatis, 0-3 septatis, non
geniculatis, apex rotundatis vel attenuatis, interdum truncatis,
cicatricibus conidialis interdum visibilis (2 um diam.), 50-100 x 7-9
um. Conidiis cylindraceis, fusiformis vel obclavato-cylindraceis,
pallide olivaceis, rectis vel leniter curvatis, laevis, frequentissime
constrictis, 1-5 septatis, apice rotundatis, basi attenuatis et
truncatis, 24-87 x 7-9 um.
Habitat in foliis vivis Oroxylt indtet Vent. (Bignoniaceae), ad
Garia, 24-Parganas, Bengal occidentalis, leg. B.K. Das, 26 XII 1979, No.
PCC 4091a (IMI 256518a) (Herb. LAM Yen #10595, holotypus).
Leaf spot indistinct, only some pale reddish-brown
angular discolored areas with indistinct margin, more or
less. vein-limited, scattered or confluent, 2-5 mm in diam-
eter, more visible on upper surface, but invisible on lower
surface: CGaespitudi strictly hypophylious, invisible weven
under hand lens. Mycelium internal: hyphae pale olivaceous,
smooth, septate, branched, 2.5-4.5 wm wide. Stromata
lacking. Conadvophoresssolitaryyor 2-3.in fascicles
emerging through the stomata, pale olivaceous and paler to-
Wards, the tip (almost hyaline), sample, straight or ?siuonery
curved... 0-5, Séptate.. not pens culace, apexyrounded.omraete rr.
ate and truncate, conidial scars sometimes visible (2 ym in
diameter), 50-100-x.7-9 jms Contdia cy lindric,. fusitouimeon
obclavate-cylindric, pale olivaceous, straight or slightly
curved, 1-5 septate, smooth, often constricted atsseptum,
apex rounded, base attenuate and subtruncate, 24-87 x 7-9
pm.
On Oroxylum indicum Vent. (Bignoniaceae), in Garia, 24-
Parganas, West Bengal), india, leo. BK. Das\ 260,471 107 0.
No. PCC 409laV (IMI 2565184), (Herbs LAM Yen #10595).
Noté:. Gercospora oroxsyli Karag Mandal differs from this
fungus by its hyaline and filiform conidia.
5. Pseudocercospora pavettae-indicae (Gov. § Thirum.)
Yen, Kar .G.Das, comb. nov. (Fig. 5, A-E) = Cerecospor:
pavettae-indtege Gov. & Thirum., Sydowia 100271, 19577
Leak spot distinct , irregularly angular, more or less
vein-limited, pale brown, sometimes without distinct margin,
but sometimes with a linear raised dark-brown margin which
is a discoloration of the limiting veinsey scattered oreo.
fluent, 1-4 mm in diameter, clearer on upper surface.
Caespituli amphigenous, visible as small dark punctiform
bodies on both surfaces. Primary mycelium internal: hyphae
pale olivaceous, smooth, branched, septate, 3-4 um wide.
Stromata well developed, black-brown, globular or subglobular,
25-50 um in diameter. Conidiophores pale olivaceous, 5-25
Fig. 4. Pseudocercospora oroxyligena: A, Young conidiophores; B
Pormation of conidia; C, 01d conidiophores; D, Conidia.
’
65
66
in fascicles emerging through the stomata, simple, straight
when young, flexuous when old, 0-3 septate, 0-2) geniculate,
sometimes with membrane finely rugose, apex rounded or
attenuate and subtruncate, conidial scars not distinct,
12-50 x 4-5 (-6) um. Secondary mycelium superficial: hyphae
pale olivaceous, arising from the stomata, séptate) branched,
3-3.5 um wide, bearing the secondary conidiophores as lateral
branches; “Conidia cylindric or obclavate=cy bindrie: pare
olivaceous, straight or slightly curved, smooth not ‘con-
stricted, 3-6 septate, apex rounded, base attenuate and
truncace, 55-50 x (3-5 66) jim.
On Pavetta indica L. (Rubiaceae), in Amtalla, 24-Parganas,
West) Bengal, India, leg. B.K. Das, 2571 X°197S), Nowe CCe ae
(IMI 242957) (Herb. LAM Yen #10596).
Note: The systematic characters of this fungus are
those of Cercospora pavettae-indicae Gov. § Thirum, which
Govindu and Thirumalachar (1957) have described. (But tor
lack of thickened conidial scars, this fungus: ought to be
transferred to genus Pseudocercospora (Ps. pavettae-indicae).
6. Pseudocercospora stillingiae (E11. G0 EvVe yen. kare
Das, comb. nov. (Fig. 5, F-1) = Cercospora stillingtae
ba weg Oe Ge OUT SMV COL wh one nel O.o gs
Leaf spot orbicular or suborbicular, deep brown, border-
ed by a yellowish zone without distinct margin, scattered,
about 2-6 mm in diameter. Caespituli amphigenous, but
chiefly hypophyllous, punctiform on lower surface of old
spot, but not visible on young ones. Primary mycelium in-
ternal: hyphae olivaceous or pale olivaceous, branched,
septate, 2-4 um wide. Stromata well developed, dark brown,
subglobular, 20-50 um in diameter. Conidiophores numerous
in dense fascicles emerging through the stomata, pale
olivaceous and very pale olivaceous towards the tip, simple,
flexuous, 0-3 septate, 0-2 geniculate, apex rounded or
shouldered, conidial scars sometimes visible, 10-30 x 3-4
um. Secondary mycelium superficial: hyphae pale olivaceous,
arising from the stomata, branched, septate, 2-4 um wide,
bearing the secondary conidiophores as lateral branches.
Conidia cylindric or obclavate-cylindric, pale olivaceous,
straight or slightly curved, smooth, not ‘constricted sus4
septate (generally 3-4 septate), ‘apex rounded, base obconic-
truncate, 14-45. 30972 5- 4) wi.
On Sapium sebiferum Roxb. (Euphorbiaceae), in Duttapurkur,
24-Parganas, West Bengal, India, lepiw BK, Dace eis .ioe
No. PCC 4061 (IMI 246496) (Herb. LAM #10598). ;
Note: This fungus shows the systematic characters of
Cercospora stillingiae Ell. § Ev. On the other hand,
Sawada (1943) has described a Cercospora sapii-sebiferi
Sawada in Taiwan, but the description and the figures show
that is a synonym of Cercospora stillingiae Ell. § Ev. But
for lack of thickened conidial Scars, this species oucicero
be transferred to genus Pseudocercospora (Ps. stillingiae).
67
=_
a
Fig. 5. Pseudocercospora pavettae-indicae(A-E): A, Primary conidio-
phores; B, Young conidiophores and fromation of Conddad: | Caan Diy oCCon]
dary conidiophores; E, Conidia.----Pseudocercospora stillingiae (P=) :
F,Young conidiophores; C,0ld conidiophores; H,Secondary conidiophores;
Tae Convddias
68
7. Pseudocercospora tectonicola Yen; Kang pac op.
nove (Pio. 6)
Maculis indistinctis, Caespitulis hypophyllis, effusis atro-
brunneis, dense velutinis. Mycelium immersum: hyphis pallide olivaceis,
laevis, septatis, ramosis, 2-4 um latis. Stromatibus plerumque nullis,
interdum mediocriter evolutis, atrobrunneis, subglobosis usque ad 20 ym
diam. Conidiophoris hypophyllis, 2-12 in fasciculo per stomatibus
emergentis, interdum solitariis, valde brunneis, sursum pallidioris,
Simplicibus, 0-12 septatis, non geniculatis, inferne erectis et laevis,
superne leniter flexuosis et rugosis, ad apicem rotundatis vel sub-
cOniCIsS, Cicatricibus conidialis indistinctis, S$6=120 x 525-2 im:
Conidiis obclavatis vel obclavato-cylindraceis, profunde brunneis,
plerumque curvatis, raro rectis, 3-9 septatis, laevis, leniter con-
strictis, apice rotundatis, basi attenuatis et truncatis, 30-100 x G.5-
8 um.
Habitat in foliis vivis Teetonae grandts L. (Verbenaceae), ad
Santipur, Nadia, Bengal occidentalis, India, leg: Bik. Bas, 25) X01 1970.
No. PCC 4102 (IMI 246495) (Herb. LAM Yen #10597, holotypus).
Leaf spot indistinct or none. Caespituli hypophyl louse
deep dark brown, effuse, densely velutinous. Mycelium in-
ternal:» hyphae pale olivaceious, smooth, septate, branched,
2-4 um wide. Stromata generally absent, sometimes poorly
developed, dark-brown, subglobular, 20 wm in diameter. Con-
idiophores hypogenous, 2-12 in fascicles emerging through
the stomata, sometimes solitary, deep-brown, paler towards
the tip, simple, straight when young, slightly flexuous when
old, 0-12 septate, not geniculate, smooth below, rugose and
more or less undulated above, apex rounded or subconic and
sometimes swollen, conidial scars not distinct, 36-1705
2.0-6 ym. CGonidia obclavate or obclavate-cylindiric, deep-
brown, generally slightly curved, sometimes straight, 3-9
septate, smooth; ‘sometimes with constrictions at the septum
apex rounded, base attenuate-truncate, 30-100 x 6.5-8 um.
On Pectona grandis L. (Verbenaceae), an Santipur, Nadeae
West Bengal, “India, legs B.K) Das, 28 XIV 1079. Now PCC
(IMI 246495) (Herb. LAM Yen #10597).
Note: The Cercospora tectonae Stevens differs from
this fungus in having hyaline conidia.
8. Pseudocercospora viticigena Yen, Kar & Das, sp. nov.
GEIR 77)
Maculis aliquantum distinctis, atro-brunneis, in epiphyllo plus_
visibilis, suborbicularis, angularis vel irregularis, dispersis, 1-3 mm
diam., interdum usque ad 4 mm longis. Caespitulis hypophyllis, invisi-
bilis. Mycelium primarium immersum: hyphis subhyalinis vel pallidissime
Olivaceis, laevis, septatis, ramosis,.1.5-3.5. um latis. Stromatibus
atro-brunneis, irregulariter globosis, 24-40 um diam. Conidiophoris
districte hypophyllis, numerosis in dense fasciculatis, pallide oliva-
ceis et sursum pallidioris, simplicibus, 0-3 septatis, 0-2 geniculatis,
saepe flexuosis, in superne interdum denticulatis, ad apicem attenuatis
vel irregulariter rotundatis, cicatricibus conidialis interdum visibilis,
15-43 x 3-4 (-5) um. Mycelium secundarium sperficiale: hyphis ex
stomatibus oriundis, pallidissime olivaceis, laevis, septatis, ramosis,
2.5-3 um latis, conidiophoris secundariis lateraliter gerentis. Con-
69
Fig. 6. Pseudocercospora tectonicola:
B, Solitary conidiophores; C, Formation of conidia; D, Above part of
old conidiophores; E, Conidia.
A,Fascicles of conidiophores;
70
idiis obclavato-cylindraceis, pallide olivaceis, rectis vel leniter
curvatis, 1-6 septatis (plerumque 3 septatis), apice rotundatis, basi
obconico-truncatis, 15-65 x 2-4 um.
Habitat in foliis vivis Vitiets negundo L. (Verbenaceae), ad
Duttapukur, 24-Parganas, Bengal occidentalis, India, leg. B.K. Das, 1
III 1980, No. PCC 4256 (IMI 250374) (Herb. LAM Yen #10600, holotypus).
Leaf spot.rather distinct, dark-brown, more: visible lon
upper surface, suborbicular, angular or irregular; scarecr.
ed, 1-3 mm in diameter, sometimes up to 4 mm long. Caespi-
tuli hypogenous, invisible even under the hand lens) @?ri-
mary mycelium internal: hyphae subhyaline or very pale
olivaceous, smooth, septate, branched, 1.53.5 Gm wide.
Stromata black-brown, irregularly globular, 24-40 um in
diameter. Conidiophores strictly hypophyllous, numerous
in dense fascicles, pale olivaceous and paler towards the
tip, simple, 0-3 septate, 0-2 geniculate, often tortuous,
sometimes denticulate above, apex attenuate or irregularly
rounded, conidial scars sometimes visible, 15-43 x 3-4 (-5)
um. Secondary mycelium superficial: hyphae arising from
stomata or from stromata and mixed with the primary conidio-
phores, pale olivaceous, septate, branched, smooth, 2.5-3
um wide, bearing secondary conidiophores as lateral branches.
Conidia obclavate-cylindric, pale olivaceous, straight or
slightly curved, 1-6 septate but generally 3 septate, apex
rounded, smooth, not constricted, base obconically truncate,
15-65 x 2-4 um.
On vitex negundo |L. (Verbenaceae), in Duttapukur, 24-
Parganas, West Bengal, India, leo. B. Ky Das et bee ocae
No. PCC 4256 (IMI 250374) (Herb. LAM Yen #10600).
Note: Cercospora viticis Hll. G| Ev., = webverz Cnupp
and C. agawalii Chupp apud Agarwal §& Hasija differ from our
fungus by their strictly epigenous fruiting. On the other
hand, Pseudocercospora viticicola (Yen §& Lim) Yen and P.
vitici-quinatae (Yen) Yen are distinct from this species by
their always amphiphyllous caespituli.
9.. Stenella coffeae Yen, Kar & Das, sp. nov. (ie. ce)
Maculis irregulariter orbicularis, griseis vel obscure griseis,
margine indistinctis, dispersis vel confluentis, 2-4 mm diam. Caespi-
tulis hypophyllis, invisibilis. Mycelium primarium immersum: hyphis
Olivaceis vel pallide olivaceis, laevis, septatis, ramosis, subtiliter
verruculosis, 2.5-4 ym latis. Stromatibus atrobrunneis, subglobosis,
25-50 um diam. Conidiophoris primariis 2-14 in fasciculo per stomatibus
emergentis, brunneis vel profunde brunneis, sursum pallidioris,
Simplicibus, inferne erectis vel suberectis, superne tortuosis vel
denticulatis, laevis, 4-6 septatis, non geniculatis, ad apicem rotunda-
tis et saepe undulatis vel denticulatis et cicatricibus conidialis
atrobrunneis ornatis, 65-220 x 4-5 um. Mycelium secundarium super-
ficiale: hyphis ex stomatibus oriundis, pallide brunneo-olivaceis,
laevis, septatis, ramosis, valde verruculosis, 2-3 um latis, conidio-
phoris secundariis lateraliter gerentis. Conidiis cylindraceis,
brunneis vel pallide brunneo-olivaceis, valde verruculosis, rectis vel
leniter curvatis, interdum leniter undulatis, 3-18 septatis, non con-
strictis, utrimque rotundatis, cellula basali in hilum atrobrunneun,
33-200 x 3-5 (-6) um.
71
Fig. 7. Pseudocercospora viticigena: A, Conidiophores and stromata;
B, Formation of conidia; C, Formation of secondary mycelial hyphae; D,
Secondary conidiophores; E, Conidia.
TZ
Habitat in foliis vivis Coffeae bengalensts Roxb. (Rubiaceae), ad
Baraipur, 24-Parganas, Bengal occidentalis, India, leg. B.K. Das, 6
XEI< 19793 No. PCO 4014 (IMI 246498) (Herb. LAM Yen #10592, holotypus).
Leaf spot blurred, irregularly orbicular, gray to dark
pray, without distinct margin, scattered or ‘conftivent ez
mm in diameter. Caespituli hypogenous, invisible even under
hand lens. Primary mycelium internal: hyphae olivaceous or
pale’ olivaceous, septate, branched, finely verruculose, 75.
aaim widées “Stromata dark brown, subvlobular, 2550 une
diameter. Primary conidiophores 2-14 in fascicles emerging
through the.-stomata, brown to deep brown, paler towards the
tip, simple, straight or substraipht below; very cortuou:
above, smooth, 4-16 septate, not geniculate, apex arounded
and often undulated or denticulate, decorated with thickened
conidial scars on the shoulders, 65-220 x 4-5 qm. Seconda...
mycelium superficial: hyphae pale brown-olivaceous, arising
from the stomata, septate, branched, strongly verruculose,
2-3 um wide, bearing secondary conidiophores which are much
more shorter than the primary conidiophores. Conidia
cylindric, brown. or pale brown-olivaceous, stronglyoverrucy—
lose, straight or Slachtly- curved, sometimes slightly un-
dulate, 3-18 septate, not constricted, rounded on both ends,
decorated with dark-brown cicatrice on the hilum, 33-200 x
3-5 (-6)° pm.
On Coffea bengalensis Roxb. (Rubiaceae), in Baraipur,
24-Parganas, West Bengal, India, leg;. B.KY Das, 6 XII
No. PCC 4014 (IMI 246498) (Herb. LAM Yen #10592).
Note: “According to Chupp (1953); Seymour (1967) ana
Ellis (1976), there are three species of Cercospora (Cercos-
pore wcorfetcola Berk. .& Cké., Ca herrerana’ Parn. and. c.
coffeae Zimm.) which are parasites on the leaves of Coffea.
Bue recently, Holliday (1980) considers that’ cCercospora
herrerana Farn. and C. coffeae Zimm. .are just the synonyms
ofACercospora cofteidola Berk,..& Cke. (The ase species
differs from our fungus by its smooth and hyaline conidia.
lO.+Stenelta garugqae Yen, Kar'’6 Das, sp. nev. (bre.
Maculis indistinctis. Caespitulis districte hypophyllis, effusis,
angularis et nervuli limitatis, atro-olivaceis, in maculo parvis, dis-
persis, 0.5-2 mm diam., interdum confluentis. Mycelium primarium immer-
sum: hyphis pallidissime olivaceis, laevis, septatis, ramosis, 1-2.5 um
latis. Stromatibus nullis. Conidiophoris primariis hypophyllis, 2-8 in
fasciculo per stomatibus emergentis, divergentis, olivaceis vel pallide
olivaceis, sursum pallidioris, laevis, simplicibus vel ramosis, inferne
erectis, superne leniter flexuosis, 0-3 septatis, 0-2 geniculatis, ad
apicem flectis et attenuatis, cicatricibus conidialis raro visibilis,
12-45 x 4-5 wm. Mycelium secundarium superficiale: hyphis ex stomatibus
vel ex conidiophoris primariis oriundis, pallide olivaceis, septatis,
ramosis, subtiliter verruculosis, 1.5-4 um latis, conidiophoris second-
ariis lateraliter gerentis. Conidiis obclavato-cylindraceis, pallide
olivaceis, rectis vel leniter curvatis, subtiliter verruculosis, non
constrictis, 3-9 septatis, apice rotundatis vel subobtusis, basi obcon-
icis et attenuato-truncatis, 32-90 x 3-4 unm.
vie)
Wick
Fig. 8. Stenella coffeae: A, Fascicles of conidiophores; B, Old
conidiophores; C, Secondary conidiophores; D, Above part of conidio-
phores; E, Conidia.
74
Habitat in foliis vivis Gargugae pitnnatae Roxb. (Burseraceae), ad
Simurali, Nadia, Bengal occidentalis, India, leg. B.K. Das, 4 XII 1980,
No. PCC 4462 (Herb. LAM Yen #10602, holotypus).
Leaf spot indistinct or none. Caespituli always hypo-
phyllous, forming olivaceous to dark olivaceous angular small
blotches on lower surface, clearly vein-limited, effuse,
scattered, 0.5-2 mm in diameter, sometimes confluent. Pri-
mary mycelium internal: hyphae very pale olivaceous, smooth,
septate, branched, 1-2.5 wm wide. Stromata absent. Conidio-
phores 2-8 in fascicles emerging through the stomata, but at
the same time secondary conidiophores arising abundantly from
secondary mycelial hyphae as lateral branches, both kinds of
conidiophores similar to each other, olivaceous or mid to
pale olivaceous and paler towards the tip, generally simple,
rarely once branched, straight below and slightly flexuous
above, 0-3 septate, 0-2 geniculate, apex often shouldered
and attenuate-truncate, conidial scars sometimes distinct,
12-45 x 4-5 wm... Secondary mycelium superficial: “hyphae
pale olivaceous, arising from the stomata or from the pro-
longation of primary conidiophores, septate, branched, very
finely verruculose, 1.5-4 ym wide, bearing abundant second-
ary conidiophores as lateral branches. Conidia obclavate-
cylindric, pale olivaceous, straischt or very slightly curved,
very finely verruculose, 3-9 septate, apex rounded’ corpse
obtuse, base obconic and attenuate-truncate, 32-90 x 3-4 um.
On Gargua pinnata Roxb. (Burseraceae), in Simurali,
Nadia, West Bengal, India; legs BiRe Das, 4X11 T9800 [3 No.
PCC 4462 (Herb. LAM Yen #10602) .
11. Stenella oroxylicola Yen, Kar & Das, sp. nov: (Fire.
10)
Maculis indistinctis, tantum area parva brunnea, margine indis-
tinctis, leniter nervuli limitatis, angularis, dispersis, 2-5 mm dian.,
interdum confluentis, in superiore superficie plus distinctis.
Caespitulis hypophyllis, effusis, in inferiore superficie griseis.
Mycelium immersum: hyphis pallidissime olivaceis, laevis, septatis,
ramosis, 2-5 ym latis. Stromatibus nullis. Conidiophoris 2-8 in fasci-
culo per stomatibus emergentis, brunneo-olivaceis, sursum pallidioris,
plerumque simplicibus, raro ramosis, laevis (cum membrana incrassatula),
saepe flexuosis, 2-13 septatis, leniter multigeniculatis, ad apicem
rotundatis vel flectis, cicatricibus conidialis interdum visibilis, 80-
260 x 6-7 um. Conidiis obclavatis, pallide olivaceis, lenitissime
curvatis, subtiliter verruculosis, non constrictis, 3-6 septatis, ad
apicem subconicis, basi attenuato-truncatis, 40-105 x 5-6 um.
Habitat in foliis vivis Oroxylt indict Vent. (Bignoniaceae), ad
Garia, 24-Parganas, Bengal occidentalis, India, leg. B.K. Das, 26 XII
1979, No. PCC 4091 (IMI 256518) (Herb. LAM Yen #10594, holotypus).
Leaf spot indistinct, only some brownish angular dis-
colored small areas without definite margin, slightly vein-
limited, more visible on upper surface, scattered, 2-5 mm
in diameter, sometimes confluent. Caespituli hypophyllous,
effuse, gray, visible on lower surface of the leaf spot.
Mycelium internal: hyphae pale olivaceous, smooth, branch-
ed, septate, 2-5 um wide. Stromata absent. Conidiophores
2-8 in fascicles emerging through the stomata, brown-
75
hores; B, Young
c, Secondary
Fascicle of conidiop
dary mycelial hyphae;
Stenella garugae: Tye
es and formation of secon
Dy Conidia.
pe pone
conidiophor
conidiophores;
76
olivaceous and paler towards the tip, generally simple.
occasionally branched, smooth with rather thick wall, almost
£lexuous, 2-13 septate, slightly multigeniculate, apex
rounded or shouldered and attenuate, conidial scars sometimes
visible, 80-260 x 6-7 um. Conidia obclavate, pale olivaceous,
slightly curved, finely verruculose, 5-0 septate, (ioc.
stricted, apex subconic, base obconic-truncate,) 40-10 sexo
um.
On Oroxylum indicum Vent. (Bignoniaceae), in Garia, 24-
Parganas, West Bengal, India, ley. Bik. Das, 26) 1 moc.
No. PCC 4091 (IMI 256518) (Herb. LAM Yen #10594).
12. Stenella xeromphigena Yen, Kar & Das, sp. nov. (Fig,
puts)
Maculis distinctis, plerumque orbicularis, in epiphyllo albo-
griseis, in hypophyllo pallide brunneis, autem margine linearis sub-
elevatis, dispersis, 3-14 mm diam., interdum confluentis. Caespitulis
amphiphyllis, autem principaliter epiphyllis, in superiore superficie
obscure punctiformis. Mycelium primarium immersum: hyphis olivaceis,
frequentissime subter cuticulis repentis, septatis, ramosis, 2.5-3 um
latis. Stromatibus subter cuticulis positis, atrobrunneis, globosis
vel subglobosis, 30-55 ym diam. Conidiophoris primariis 12-42 in
fasciculo per scissuris cuticulis emergentis, olivaceis, concoloris,
laevis, simplicibus, erectis, cylindraceis, 1-3 septatis, non genicula-
tis, ad apicem irregulariter rotundatis vel attenuatis, cum cicatricibus
conidialis atrobrunneis ornatis, 14-65 x 3-4 um. Mycelium secundarium
superficiale: hyphis olivaceis vel pallide olivaceis, hypophyllis,
ex stomatibus oriundis, repentis vel arcuatis, subtiliter verruculosis,
septatis, ramosis, 2.5-3.5 um latis, conidiophoris secundariis lateral-
iter gerentis. Conidiis cylindraceis vel vermiformis, olivaceis,
rectis vel leniter curvatis, interdum undulatis, solitariis vel catena-
tis, utrimque rotundatis, interdum utrimque cicatricibus atrobrunneis
ornatis, plerumque cellulis basali in hilum cicatricibus atrobrunneis
ornatis, 20-156 x 3-4 un.
Habitat in foliis vivis Xeromphidts ultginosae (Rubiaceae), ad
Sylva Raiganj, Dinajpur occidentalis, Bengal occidentalis, India, leg.
B.K. Das, 25 III 1980, No. PCC 4286 (IMI 250390) (Herb. LAM Yen #10593;
holotypus).
Leaf spot distinct, generally orbicular, sray-whitish
on upper surface, pale brownish on lower surface, but with
a linear raised margin on both surfaces, scattered, 3-14-mm
in diameter, sometimes confluent. Caespituli amphiphyllous,
but chiefly. epiphyivous, visible as small dark, punctirerm
on upper surface of the leaf spot. Primary mycelium internad,
hyphae olivaceous, often under the cuticle, septate, branc’
ed, 2.5-3.5 wm wide. Stromata well developed, generally
Situated between the cuticle and the epidermal cells, dark-
brown, globular or subglobular, 30-55 wm in diameter. Pri-
mary conidiophores 12-42 in fascicles emerging through the
TUpcure Of -cutrelé, olivaceous. and concolorous, simple,
cylindric and straight, 1-3 septate, apex irregularly
rounded or attenuate-truncate and decorated with dark-brown
conidial scars, 14-65 x 3-4 um. Secondary mycelium super-
ficial: hyphae olivaceous or pale olivaceous, hypophyllous,
abundant, arising from the stomata, repent or arcuate,
finely verruculose, branched, septate, 2.5-3.5 um wide,
Ta
Fig. 10. Stenella oroxylicola:A, Old conidiophores; B, Young
conidiophores; C, Branched conidiophores; D, Formation of conidia;
Bp Conidia.
78
Fig. 11. Stenella xeromphigena: A, Conidiophores and stromata;
B, Young conidiophores; C, Old conidiophores; D, Secondary conidio-
phores; E, Conidia.
ib!)
bearing secondary conidiophores as lateral branches. Conid-
ia cylindric-or narrowly vermiform, olivaceous, straight or
curved and undulated, finely verruculose, 1-14 septate, not
constricted, solitary or catenate, rounded on both ends,
generally decorated with a dark-brown cicatrice on the hilun,
but sometimes both ends decorated by two dark-brown cica-
ErIcesn ac 150 xX 3-4 jm,
On leaves of Xeromphis uliginosa (Rubiaceae), in Raigani
Forest, West Dinajpur, West Bengal, India, leg. B.K. Das,
Zot tos No. PCC) 4286 "CIM! 250390) ’(Herb EAM. Yen
#10593).
ACKNOWLEDGEMENT
We are grateful to Dr. F. C. Deighton and Commonwealth Mycological
Institute, Kew, England, for the help given in the determination of
critical specimens. We are much indebted to Dr. Richard P. Korf,
Professor of Mycology of Cornell University and Dr. Don R. Reynolds,
Curator in Botany of Los Angeles County Natural History Museum, for re-
viewing the manuscript.
DELLE RATLURE Ci hE D
Boedijn, K. B. 1961. The genus Cercospora in Indonesia. Nova Hedwigia.
3:424,
Ciipp, Cy 19530 A) monograph of the genus Cercospora., pp. 212, 0231,
SOT » AIS eee ley po76:, ‘anda o 7;
Deighton, F. C. 1959. Studies on Cercospora and allied genera 1. Cer-
cospora species with coloured spores on Phyllanthus (Euphorbiaceae).
Mycol. Pap. 71:3-23.
Deighton, F. C.. 1976. Studies on Cercospora and allied genera VI.
Pseudocercospora Speg., Pantospora Cif., and Cercoseptoria Petr.
Mycol, Pap. 140:40, 138, and 145.
Ellis, M. B. 1976. More-Dematiaceous Hyphomycetes. p. 283.
Govindu, H. C. & Thirumalachar, M. J. 1957. Notes on some Indian
Cercosporae. Sydowia, 10:271.
Holliday, P. 1980. Fungus diseases of tropical crops. p. 68.
Kar, A. K. & Mandal, M. 1969. New Cercospora spp. from West Bengal.
Trans. Br. Mycol, Soc., 55:344;
Pavgi, M. S. & Singh, M. P. 1966. Parasitic fungi from North India
VII. Mycopath. Mycol. appl. 30:267.
Sawada, Ke 1921. "Descr.. Cat, Formosan Fungi, 2:153.
Sawada, K. 1943. Descr. Cat. Formosan Fungi, 8:99.
Sawada, K. 1959, Descr. Cat. Formosan Fungi, 11:227.
Seymour, A. B. 1967. Host index of the fungi of North America. p. 618.
Yen, J. M. 1977. Etudes sur les champignons parasites du Sud-Est
asiatique 26. Les Cercospora de Formose II.! Bull. Soc. Mycol.
Fro poo doo.
Yen, J. M. 1978. Etudes sur les champignons parasites du Sud-Est
aSsiatique 33. Les Cercospora de Formose V. Pseudocercospora.
Bull Soc. Mycol. Fr.-94:385-389.
Yen, J. M. & Lim, G. 1973. Etudes sur les champignons parasites du
Sud-Est asiatique 20. Dixieme note sur les Cercospora de Malaisie.
Cahiers du Pacifique, No. 17:104.
Yen, J. M. & Lim, G. 1980. Cercospora and allied genera of Singapore
and the Malay Peninsula. Gardens' Bulletin, Singapore. 33:190.
MYCOTAXON
Vol exV ie Nos wi epp. OU o'5 October-December 1982
STUDIES ON HYPHOMYCETES FROM WEST BENGAL, INDIA, III.
CERCOSPORA AND ALLIED GENERA OF WEST BENGAL, 3
JO-MIN YEN
Natural History Museum, 900 Exposition Boulevard
Los Angeles, California 90007, U.S.A.
and
A. Ke. KAR and B.~ kK. DAS
Department of Botany, Presidency College, Calcutta-700073, India
ABSTRACT
The third of a series of studies on Hyphomycetes from
West Bengal, India, includes descriptions and illustrations
of six new species of Indian Cercosporae: Cercoseptorta
ee: sp. nov., on Caesalpinta digyna; C. cedrelae
sp. nov., on Cedrela toona; Phaeotsartopsis caesalpintae
Sp. nov., on Caesalpinta bonducella; Pseudocercospora melt-
teola sp. nov., on Melta azedarach; Stenella cedrelae sp.
nov., on Cedrela toona; and Stenella cynancht sp. nov., on
Cynanchum callttata.
1. Cercoseptoria caesalpiniae Yen, Kar @ Das, sp, nov.
(CBig® 1)
Maculis indistinctis. Caespitulis districte hypophyllis, invisi-
bilis. Mycelium immersum: hyphis pallidissime olivaceis vel sub-
hyalinis, laevis, septatis, ramosis, 2.5-5 um latis, interdum usque ad
7 um latis, substomatalis. Stromatibus nullis vel valde inchoatis.
Conidiophoris hypophyllis, 2-15 in fasciculo per stomatibus emergentis,
pallide olivaceis, concoloris, simplicibus vel ramosis, erectis vel
leniter flexuosis, 1-6 septatis, 0-1 geniculatis, ad apicem rotundatis,
cicatricibus conidialis invisibilis, 30-60 x 4-5 um. Conidiis cylin-
draceis, pallide olivaceis, rectis vel leniter curvatis, plerumque 3
septatis, interdum 5 vel 7 septatis, laevis, non constrictis, apice
rotundatis, basi truncatis, 35-85 x 3-3.5 um.
Habitat in foliis vivis Caesalpinitae digynae Rottb. (Leguminosae),
ad Aranghata, Nadia, Bengal occidentalis, India, leg. B.K. Das, 29 XII
1979, No. PCC 3708 (IMI 237401) (Herb. LAM Yen #10610, holotypus).
Leaf spot indistinct or none. Caespituli strictly hypo-
phyllous, invisible even under the hand lens. Mycelium
internal: hyphae very pale olivaceous or subhyaline,
smooth, branched, septate, 2.5-5 wm wide, up to 7 um for
thatihsituated beneath the stomata. Stromata Jacking omvery
rudimentary. Conidiophores hypophyllous, 2-15 in fascicles
81
Fig. 1. Cercoseptoria caesalpiniae: A, Branched conidiophores; B,
Young conidiophores and rudimentary stromata; C, Formation of
conidia; D, 3 septate conidia; E, 5-7 septate conidia.
82
emerging through the stomata, pale olivaceous, concolorous,
simple or branched, straight or slightly flexuous, 1-6
septate, 0-1 geniculate, smooth, apex rounded or attenuate
and sometimes slightly swollen, conidial scars indistinct,
30-60%x 4-50pm.. Conidia cylindric; pate oltvaceous,
straight or slightly curved, generally 3 septate (sometimes
5 or 7/,.septate), ‘smooth, not constricted, apex. 1ounded,
base truncate, (35-65 x, 5-5..5 um.
On living leaves of Caesalpinia dtgyna Rottb. (Legumi-
nosaeée),iiin Aranghata, Nadia, West Bengal, India, leg. ye ox
Das, 29 XIT 1979, No. PCC 3708 (IMI 257401)"9 (Hero. Ameren)
#10610).
2. Cercoseptoria cedrelae Yen, Kar §& Das, sp. nov. (Fig. 2)
Maculis angularis vel irregularis, saepe nervuli limitalis, cum
margine aliquantum distinctis, primo lucido-brunneis, dein in epi-
phyllo albo-griseis et in hypophyllo obscure brunneis, dispersis, in-
terdum confluentis, 1-8 mm diam. Caespitulis amphigenis, autem princi-
paliter hypophyllis, atro-punctiformis. Mycelium primarium immersun:
hyphis pallidissime olivaceis, laevis, septatis, ramosis, 2-2.5 um
latis. Stromatibus atro-brunneis, globosis vel subglobosis, saepe
intra cellulis epidermicis, 20-30 um diam. Conidiophoris 5-32 in
fasciculo per scissuris cellulis epidermicis emergentis, pallide
Olivaceis vel olivaceis, concoloris, simplicibus, laevis, 0-1 septatis,
O-1 geniculatis, cicatricibus conidialis indistinctis, ad apicem
irregulariter rotundatis, 14-35 x 3.5-4.5 um. Mycelium secundarium
superficiale: hyphis ‘pallide olivaceis, ex stomatibus oriundis,
laevis, septatis, ramosis, 2-3 um latis, conidiophoris secundariis
lateraliter manifestibus. Conidiis filiformis, pallidissime olivaceis,
leniter curvatis, plerumque 5 septatis (raro 2 vel 6 septatis), laevis,
non constrictis, apice obtusis, basi truncatis, 50-80 x 2-2.5 um.
Habitat in foliis vivis Cedredae toonae Roxb. (Meliaceae), ad Rai-
ganj, Dinajpur occidentalis, Bengal occidentalis, India, leg. B.K. Das,
7 X 1980, No. PCC 4415a (Herb. Lam Yen #10608, holotypus) .
Leaf spot. distinct, angular’ or irregular, often, vyein-
limited, with margin-rather, distanct,, at tarst brownish
becoming later to white-grayish on upper surface and dull
brown on lower surface, scattered or confluent, 1-8 mm in
diameter. Caespituli amphigenous, but chiefly hypophyllous,
black punctiform. Primary mycelium internal: hyphae pale
olivaceous, smooth, branched, septate, 2-2.5 wm wide.
Stromata globular or subglobular, dark brown, often situ-
ated in the epidermal cells, 20-30 um in diameter. <‘Conidic-;
phores 5-32 in fascicles emerging through the rupture of the
epidermal cells, pale olivaceous to mid olivaceous, con-
colorous, simple, smooth, 0-1 (rarely 2) septate, 0-1
geniculate, conidial scars not visible, apex irregularly
rounded (sometimes shouldered), 14-35 x 3.5-4 um. Secondary
mycelium superficial: hyphae pale olivaceous, arising from
the stomata, smooth, branched, septate, 2-3 um wide, bearing
secondary conidiophores as lateral branches. Conidia very
pale iolivaceous, “filiform, sliehtly curved, ‘veenenally ms
septate (rarely 6 septate), smooth, not constricted, apex
obtuse, base truncate, 2-2.5 wm wide and 50-80.m long.
On living leaves of Cedrela toona Roxb. (Meliaceae), in
83
Fig. 2. Cercoseptoria cedrelae: A, Old conidiophores and stromata;
B & C, Young conidiophores and stromata; D, Formation of secon-
dary mycelial hyphae; E, Secondary conidiophores; F, Conidia.
84
Raiganj, West Dinajpur, West Bengal, India, tego bak Dace
7 X 19805) No. 8PCG 4415a (Herb. LAM Yen | boogie).
3, Phaeoisariopsts caesalpiniae Yen, KameG Das. icp nov.
CR. 33)
Maculis orbicularis vel suborbicularis, clare brunneis, minusculis,
1-2 mm diam., dispersis, cum marginis atro-linearis et subelevatis cir-
cumtextis, raro confluentis. Caespitulis districte hypophyllis, atro-
punctiformis. Mycelium immersum: hyphis subhyalinis, laevis, ramosis,
septatis, 2.5-3.5 um latis. Stromatibus evolutis, irregulariter globo-
sis, 30-45 um diam. Conidiophoris hypophyllis, pallide brunneo-
olivaceis, sursum pallidioris, laevis, 12-32 in fasciculo synnematiforme
aggregatis, multi-septatis, simplicibus, plerumque erectis, ad apicem
rotundatis, cicatricibus conidilis distinctis (2-2.5 um diam.) , 50-215x
4-6 um. Conidiis obclavatis vel obclavato-cylindraceis, pallidissime
olivaceis, plerumque curvatis, raro rectis, 2-9 septatis, laevis,
-apice rotundatis, basi obconico-truncatis, 43-105 x 5-6.5 um.
Habitat in foliis vivis Caesalptntae bonducellae (L.) Flem.
(Leguminosae), ad Palpara, Nadia, Bengal occidentalis, India, Leg. B.K.
Das, 25 XI 1979, No. PCC 3706 (IMI 237382) (Herb. LAM Yen #10611, holo-
typus) .
Leat spot distinct, orbicular or suborbicular, bison
brown, often bordered with a raised dark linear margin on
both two surfaces, scattered or slightly confluent, “l-2emm
in diameter. Caespituli always hypophyllous, in dark puncti-
form. Mycelium internal: hyphae subhyaline, smooth, septate,
branched, 2.5-3.5 um wide. Stromata well developed, sub-
stomatal, irregularly globular, 30-45 um an diameter.) Gon:
ophores hypophyllous, synnematous, 12-32 in dense synnemata
arising from the well developed stromata, olivaceous-brown
and paler towards the tip, simple, multiseptate, generally
straight, not geniculate, smooth, dark-brown conidial scars
Visible, at the tip of young: conidiophores (2-2) 5.7mm
diameter), apex rounded, 50-215 x 4-6 um. Conidia obclavate
or obclavate-cylindric, very pale olivaceous, generality
curved, 2-9 septate, smooth, not constricted, apex rounded,
base obconic-truncate, 43-105 x 5-6.5 um.
On living leaves of Caesalpinita bonducella (L.) Flem.
(Leguminosae), in Palpara, Nadia, West Bengal, India, leg:
Ban. Das, 2 XL 1979.,No, PCC 3706 (IML 237382) (Herb on
Yen #10611).
4. Pseudocercospora kashotoensts (Yamamoto) Deighton (Fig. 4)
CMI Mycological. Papers, No., 140.1462 [007 Gr.
= Cercospora kashotoensts Yamamoto, Trans. Nat. Hist. Soc.
FPOUVMOS a. 20.2527, 919.50),
Leaf spot indistinct or none. Caespituli hypophyllous,
slightly effuse, forming pale olivaceous blotches on lower
surface, without any boundary. Primary mycelium internal:
hyphae very pale olivaceous or subhyaline, smooth, septate,
branched, 2-3 yum wide. Stromata none. Conidiophores hypo-
phyllous, 2-7 in: loose fascicles emerging through the sto-
mata, brown-olivaceous or pale brown, simple or rarely
branched, straight and attenuate when young, flexuous or
85
Fig. 3. Phaeoisariopsis caesalpiniae: A, Stromata; B, Young conidio-
phores: C, Synnemata(caespitose conidiophores); D, Formation
Ge Comukeliiep i, Comsrclile-
86
sinuous when old, 1-4 septate, 0-2 geniculate, sometimes
forming pseudo-annellations, apex rounded or shouldered,
conidial scars scarcely visible but sometimes visible on
the shoulders or at the end of short denticles (1.5-2 jm in
diameter), 25-75 x 4-6 um. Secondary mycelium superficial:
hyphae pale olivaceous, arising from the stomata, smooth,
septate, branched, 2-3 um wide, bearing secondary conidio-
phores as lateral branches. Conidia cylindric or obclavate-
cylindric, very pale olivaceous, straight or slightly ‘curved,
1-7 septate, apex subacute to subrounded, base attentuate-
truncate of obconic-trincate; 25-105 kos oem:
On living leaves of Clerodendron itnerme (L.) Gaertn.
(Verbenaceae), in Andul, Howrah, West Bengal, India, leg.
B.K. Das, 20 XII 1979, No. PCC 4082 (IMI 254413) a( ferro. aw
Yen #10604).
Note: The Pseudocercospora clerodendrt (Miyake) Deigh-
ton differs from this fungus by its distinct suborb reulae
to angular leaf spot and especially by its fruiting amphi.
phyllous or almost epiphyllous.
Distribution: Taiwan (China) and West Bengal (India).
5. Pseudocercospora meliicola Yen, Kar § Das, sp. nov. (Fig.
5)
Maculis distinctis, in epiphyllo plus visibilis, angularis vel
irregularis, per nervuli limitatis, in inferiore superficie lucido-
brunneis, in superiore superficie albo-brunneis, dispersis vel leniter
confluentis, plerumque 1-4 mm diam., interdum usque ad 8 x 4 mn.
Caespitulis amphigenis, autem principaliter epiphyllis. Mycelium
primarium immersum: hyphis pallide olivaceis, laevis, ramosis, septatis,
2-3 um latis. Stromatibus atrobrunneis, subglobosis, 25-35 um diam.
Conidiophoris amphiphyllis, numerosis in fasciculo per stomatibus
emergentis, pallide brunneo-olivaceis, concoloris, plerumque simplicibus,
interdum ramosis, flexuosis, 0-3 septatis, 0-2 geniculatis, laevis, ad
apicem rotundatis vel attenuatis, cicatricibus conidialis indistinctis,
15-45 (-50) x 3-4.5 um. Mycelium secundarium superficiale: hyphis
pallide olivaceis, cum conidiophoris primariis commixa in fasciculo
per stomatibus emergentis, septatis, ramosis, laevis, 2-3.5 um latis,
conidiophoris secundariis lateraliter gerentis. Conidiis obclavato-
cylindraceis, pallide olivaceis, plerumque leniter curvatis, 3-8 (-11)
septatis, apice rotundatis vel subobtusis, basi attenuatis vel obconico-
truncatis, 28-92 (-120) x 3-4 um.
Habitat in foliis vivis Meltae azedarach L. (Meliaceae), ad Mallick-
pur, 24-Parganas, Bengal occidentalis, India, leg. B.K. Das, 4 IX 1980,
No. PCC 4352 (Herb. LAM Yen #10609, holotypus).
Léaf spot distinct, much more visible von upper sugiace.
angular or irregular, vein-limited, brown-whitish on upper
surface and bright brown on lower surface, scattered or
slightly confluent, 1-4 mm in diameter, sometimes up to 8 x
4 mm. Caespituli amphigenous, but chiefly epiphyllous.
Primary mycelium internal: hyphae pale olivaceous, smooth,
branched, septate, 2-3 um wide. Stromata dark-brown, sub-
globular, 25-35 um in diameter. Conidiophores numerous in
fascicles emerging through the stomata, pale brown-olivaceous,
concolorous, generally simple, sometimes branched, flexuous,
0-3 septate, 0-2 geniculate, smooth, apex rounded or attenuates
ares
diophores; D, Pseudo-annellations; E,
87
Pseudocercospora kashotoensis: A, Young conidiophores and
formation of conidia; B, Old conidiophores; C, Branched coni-
Secondary conidiophores;
Eye COnudaa:.
88
Fig. 5. Pseudocercospora meliicola: A, Stromata and young conidio-
phores;B, Branched conidiophores; C, Old conidiophores; D, Forma-
tion of secondary mycelial hyphae; E,Secondary conidiophores; F,
Conidia.
89
CChutat IsCars indistinct, US745. (-50)yx) 3-4.5 qm. Secondary
mycelium superficial: hyphae pale olivaceous, arising from
the stomata and mixed with the primary conidiophores, septate,
branched, smooth, 2-3.5 wm wide, bearing secondary conidio-
phores as lateral branches. Conidia obclavate-cylindric,
pale olivaceous, generally slightly curved, 3-8 (-ll) septate,
apex rounded or subobtuse, base attenuate or obconically
truncate, 28-92 (-120) x 3-4 ym.
On living leaves of Melita azedarach L. (Meliaceae), in
Mallickpur, 24-Parganas, West Bengal, India, leg. B.K. Das,
Ai NeOs80 5 No, PGG-43552" (Herb. LAM Yen #10609))°.
Note: The Pseudocercospora subsessilis (H. §& P. Sydow)
Perenhtousditters trom this fungus. by) its icircular and not
vein-limited leaf spot, by its hypophyllous caespituli and
especially by its conidiophores not septate, not branched
and not geniculate.
6. Pseudocercospora phyllanthi-ntrurt (Yen) Yen (Fig. 6, D&E)
Gardens oulletin Singapore 33: 181, 19380.
= Cereospora phyllanthi-nirurt Yen, Revue de Mycologie 32:
POZ VOGT;
Syn. Cercospora phyllanthticola Yen, REvue de Mycologie. 30:
Pepe 90 5,.
non Cereospora phyllanthteola Shakil §& Kamal, Indian Phyto-
DabNOLogy 152296, 1962.
Leat spot indistinct ‘or none. ‘Caespituli always hypo-
phyllous, etfuse, velutinous, olivaceous-gray, often extend-
iiRewand covering the whole Lower surface of the leaf:
Mycelium anternal: hyphae pale olivaceous, smooth, septate,
branched, 2.5-4 ym wide. Stromata lacking. Conidiophores
2-8 in fascicles emerging through the stomata, pale to mid
brown-olivaceous, simple and straight when young, flexuous
and branched when old,\1-7. septate, 0-2 geniculate, conidial
scars rarely visible, apex rounded, 18-110 x 4-6 ym. Conidia
obclavate-cylindric but always crescent-shaped, pale oliva-
ceous, 3 septate, apex rounded, base attenuate and obconical-
VY oruncace (52-5) “x 455-6. ms
On living leaves of Phyllanthus nirurt L. (Euphorbiaceae),
invoimurali, Nadia, West Bengal, India, leg. B.K, Das, 4 XII
boS0e Noe ePCC 4465. CIMT 254415) (Herb. LAM Yen..#10006)..
Note: This fungus differs from other species on Phyllan-
thus by its conidia always 3 septate and 4.5-6 ym wide.
Distribution; singapore: and India.
7. Pseudocercospora tremattcola (Yen) Deighton (Fig. 6, A-C)
CMI Mycological Papers 140:154, 1976.
=~cercospora tremattcord Ven, Bull. (soc. Mycol, Pr. .36:
(aes als AUR
Leaiy spot indistinct,.or none.» Caespitnl se cenerdal ly,
hypophyllous, effuse, velutinous, forming angular and vein-
limited areas,,.small, dark-gray, scattered, 1-1.5 mm in diam-
eter, sometimes confluent. Mycelium internal: hyphae oliva-
ceous, smooth, septate, branched, 3-5.5 ym wide. Stromata
none. Conidiophores 2-15 in fascicles emerging through the
stomata, pale olivaceous to mid olivaceous, simple or branch-
ed, flexuous or undulated, 3-10 septate, smooth, not genicu-
late, apex rounded or attenuate, conidial scars indistinct,
90
30-135 x 4.5-6 um. Secondary mycelium superficial: hyphae
pale olivaceous’,, arising from; the base of thevfascicies mr
primary conidiophores, septate, branched, smooth, 2-3.5 yum
wide, bearing numerous secondary conidiophores as lateral
branches. Conidia cylindric, pale olivaceous. generalhy,
straight, sometimes slightly curved, 3-10 septate, apex
rounded, base obconically truncate, 30-130 x 4.5-6 um.
On living leaves of Trema orientalis Bl. (Ulmaceae),
in Simurali, Nadia, West Bengal, India, leg. B.K. Das, 4 XII
1980, No. PCC 4464 (IMI 254414) (Herb... LAM Yem #10612).
Distribution: Singapore, Taiwan (China) and India.
8. Pseudocercospora tritumfettae (H. Sydow) Deighton
CMin Mycological Papers 1402122) 19.76:.
=Cercospora triumfettae H. Sydow, Ann. Mycol. 28:218,
1930.
On living leaves of Triumfetta rhombotdea L. (Tillia-
ceae), in Hemtabad Forest, West Dinajpur, West Bengal, India,
leg. BK. Dasiii7 X°1980 . Now PCC 4403 CIM ot 40s tierra
LAM Yen #10605).
Note: We have sent the materials of this fungus to
Commonwealth Mycological Institute, Kew, England, and it is
identified as Pseudocercospora triumfettae (H. Sydow) Deigh-
ton. Deighton (1976) has well described and illustrated all
the, characters or this species.
Distribution: Venezuela, Brazil, Bermuda, Cuba keep.
Dominica, and India.
Oo. Stenella cedreLae: Yen, Kar § Das,..sp. nov. .(iam
Maculis distinctis, angularis vel irregularis, denique in hypo-
phyllo obscure brunneis, in epiphyllo albo-griseis, dispersis, 1-8 mm
diam., interdum confluentis. Caespitulis amphigenis, autem principal-
iter hypophyllis, atro-punctiformis. Mycelium primarium immersum:
hyphis pallide olivaceis, laevis, septatis, ramosis, 2-3 ym latis.
Stromatibus nullis. Conidiophoris 2-25 in fasciculo per stomatibus
emergentis, simplicibus vel ramosis, pallide olivaceis, concoloris,
laevis, flexuosis, 1-3 septatis, 0-2 geniculatis, ad apicem irregular-
iter rotundatis, cicatricibus conidialis invisibilis, 22-50 x 4-5 um.
Mycelium secundarium superficiale: hyphis pallide olivaceis, subti-
liter verruculosis, ex stomatibus oriundis, ramosis, septatis, 2-3.5
um latis, conidiophoris secundariis lateraliter gerentis. Conidiis
obclavatis vel obclavato-cylindraceis, plerumque leniter curvatis, sub-
tiliter verruculosis, non constrictis, 5-15 septatis, apice obtusis
vel subrotundatis, basi obconico-truncatis, 42-160 x 3-4.5 um.
Habitat in foliis vivis Cedrelae toonae Roxb. (Meliaceae), ad
Raiganj, Dinajpur occidentalis, Bengal occidentalis, India, leg. B.K.
Das, 7 X 1980, No. PCC 4415 (Herb. LAM Yen #10608, holotypus).
Leaf spot distinct, angular or irregular, often vein-
limited, margin rather distinct, at first small ane Drags
brown, then becoming dull brown on lower surface and whitish-
gray on upper surface, scattered, sometimes confluent, 1-8
mm in diameter. Caespituli amphigenous, but chiefly hypo-
phyllous, in black punctiform. Primary mycelium internal:
hyphae pale olivaceous, branched, septate, smooth, 2-3 yum
wide. Stromata none. Primary conidiophores 2-25 in fascicl
Fig.
oF
6. Pseudocercospora phyllanthi-niruri(D & E): D, Conidiophores;
E, Conidia. ---- Pseudocercospora trematicola(A-C): A, Conidio-
phores; B, Secondary mycelial hyphae and secondary conidiophores;
C, .coniaza.
IE
emerging through the stomata, pale olivaceous, concolorous,
simple*or branched, smooth, flexuous,- 1-37 seperate
geniculate, apex irregularly rounded or shouldered, conidial
scars not visible, 22-50 x 4-5 um. Secondary myceliumesuper.
ficial: hyphae pale olivaceous, finely verruculose (sarisang
from the stomata, branched, septate, Z-3.5 jm wide, pegeme
abundant secondary conidiophores as lateral branches.
Conidia obclavate to obclavate-cylindric, generally slightly
curved, finely verruculose, not constricted, 5-14 septate,
apex obtuse or subrounded, base obconic-truncate, 42-160 x
3-4.5 um.
On living leaves of Cedrela toona Roxb. (Meliaceae), in
Raiganj, West Dinajpur, West Bengal, India; leg. BOK.) Das,
1k L980 No. PCC 4415 (Herb... LAM Yen 7 10603).
10 Sténela cynanch? Yen; Kar & Das, ‘speinov. (rrr)
Maculis indistinctis. Caespitulis hypophyllis, effusis, velutinis,
griseo-olivaceis, in inferiore superficie saepe fere totam folii pagi-
nam omnino obtengentis. Mycelium primarium immersum: hyphis olivaceis,
subtiliter verruculosis, ramosis, septatis, 2-4 um latis. Stromatibus
nullis. Conidiophoris semper ex mycelium secundarium oriundis, simpli-
cibus, pallide brunneo-olivaceis vel obscure brunneo-olivaceis, inferne
plerumque erectis et superne leniter flexuosis vel denticulatis, 1-7
septatis, non geniculatis, laevis, apex attenuatis vel denticulatis et
cicatricibus conidialis atro-brunneis decoratis, 15-135 x 3-3.5 um.
Mycelium secundarium superficiale: hyphis pallide olivaceis vel medio-
olivaceis, ex stomatibus oriundis, acute verruculosis vel echinulatis,
septatis, ramosis, 1.5-2 um latis, conidiophoris secundariis numerosis
lateraliter manifestibus. Conidiis cylindraceis vel fusiformis, pallide
olivaceis, solitariis vel catenulatis, rectis, subtiliter verruculosis,
0-1 septatis, apice rotundatis, basi semitruncatis et cicatricibus coni-
dialis atro-brunneis decoratis, interdum utrimque cicatricibus atro-
brunneis ornatis, 7-25: x 2.5-3,5 pm:
Habitat in foliis vivis Cynancht calltatatae Ham. (Asclepiadaceae) ,
ad Sylva Joyanti, Alipurduar, Jalpaiguri, Bengal occidentalis, India,
leg. B.K. Das, 21 X 1980, No. PCC 4424 (Herb. LAM Yen #10613, holo-
typus).
Leaf spot indistinct or none. Caespituli hypophyllous,
effuse, velutinous, gray olivaceous, extending and covering
the whole lower surface of the leaf. Primary mycelium
internal: hyphae olivaceous, finely verruculose, branched,
septate, 2-4 ym wide. Stromata lacking. Conidiophores
always arising from the secondary mycelial hyphae as lateral
branches, solitary, simple, pale brown-olivaceous to dark
brown-olivaceous, generally straight below and slightly
flexuous and denticulate above, 1-7 septate, not geniculate,
smooth, apex attenuate or denticulate and decorated with
numerous dark-brown conidial scars, 15-135 x 3-3. 5 un.
Secondary mycelium superficial: hyphae pale olivaceous to
mid olivaceous, arising from the stomata, branched, septate,
sharply verruculose or echinulate, 1.5-2 um wide, bearing
numerous secondary conidiophores as lateral branches.
Conidia cylindric or fusiform, pale olivaceous, solitary or
catenate, straight, finely verruculose, 0-1 septate, apex
rounded, base semitruncate and decorated with a dark-brown
93
Fig. 7. Stenella cedrelae: A, Young conidiophores; B, Old and bran-
ched conidiophores; C, Secondary mycelial hyphae and secondary
conidiophores; D, Secondary conidiophores; E, Conidia.
94
conidial scar, sometimes both two ends decorated with dark-
brown Contdial scars. (225 X 205-5. us
On living leaves of Cynanchum callitata Ham. (Asclepi-
adaceae), in Joyanti Forest, Alipurduar, Jalpaiguri, West
Bengal, India, tLéeg. B.K.. Das; 21 X 1980, No. PCG 437245 \(Hera,
LAM Yen #10613).
ACKNOWLEDGEMENTS
We are grateful to Dr. F, C. Deighton and the Directo
of the Commonwealth Mycological Institute, Kew, England, for
the help given in the determination of critical specimens.
We are much indebted to Dr. Richard P. Korf, Professor of
Mycology, Cornell University and Dr. Don R. Reynolds,
Curator of Botany, Los Angeles County Natural History Museum,
for reviewing the manuscript.
Fig. 8. Stenella cynanchi: A, Old conidiophores; B, Young conidio-
phores and formation of conidia; C, Germination of conidia;
D, Above part of old conidiophores; E, Conidia.
a5
LITERATURE CITED
Chowdhury, S. 1961. Notes on fungi from Assam, IV. Lloydia, 24:94-96.
Chupp, C. 1953. A monograph of the fungus genus Cercospora. Ithaca,
New York.
Deighton, F. C. 1976. Studies on Cerecospora and allied genera, VI.
Pseudocercospora Speg., Pantospora Cif., and Cercoseptorta Petr.
CMI Mycological Papers No. 140.
Deighton, F. C. 1979. Studies on Cercospora and allied genera, VII.
New species and redispositions. CMI Mycological Papers No. 144.
Shakil, A. K. & Kamal, M. 1962. A new species of Cercospora on
Phyllanthus ntrurt L. Indian Phytopathology 15:296-297,
Shakil, A. K. §& Kamal, M. 1963. Cercospora paktstantca, a new species
from Pakistan. Mycopathologia 21:112-113.
Yen, J. M. 1965. Etude sur les champignons parasites du Sud-Est
asiatique, III. Deuxiéme note sur quelques nouvelles espéces de
Cerecospora de Singapour. Revue de Mycologie 30:166-204.
Yen, J. M. 1967. Etude sur les champignons parasites du Sud-Est
asiatique VII. Quatriéme note sur quelques Cercospora et
Stenella de Singapour (Malaisie). Revue de Mycologie 32:179-202.
Yen, J. M. & Lim, G. 1980. Cercospora and allied genera of Singapore
and the Malay Peninsula. Garden's Bulletin Singapore. 33:
152-265,
MYCOTAXON
Vol XV, Now Lop. 96-98 October-December 1982
STUDIES ON PARASITIC FUNGI FROM SOUTH EAST ASIA, 45.
PARASITIC FUNGI FROM MALAYSIA, 22.
JO-MIN YEN
Natural History Museum, 900 Exposition Boulevard
Los Angeles, California 90007, U.S.A.
and
GLORIA LIM
Department of Botany, National University of Singapore
Kent Ridge Campus, Singapore 0511
Abstract
A new species, Stenella bougainvilleae Yen & Lim is
described and illustrated from type material on its host,
Bougainvillea spectabilis Willd., collected in Singapore. A
new combination, Cercoseptoria cordiicola Yen) Yen, is
mentioned.
Stenella bougainvilleae Yen & Lim, sp. nov.
Maculis distinctis, brunneis, orbicularis vel irregulariter
orbicularis, margine atro-brunneis circumtextibus, disper-
sis, 1-7mm diam. saepe confluentibus. Caespitulis hypophyll-
is, invisibilis. Mycelium primarium immersum: hyphae palli-
dissime brunneo-olivaceis, septatis, ramosis, 1.5-2.5um
latis. Stromatibus non evolutis. Conidiophoris primariis
hypophyllis, 2-8 in fasciculo per stomatibus emergentis,
simplicibus, raro ramosis, inferne erectis, antice atten-
uatis et denticulatis, pallide brunneo-olivaceis, sursum
pallidioribus, 3-8 septatis, O-3 geniculatis, apicibus ang-
ustioribus et cicatricibus sporarum atro-brunneis ornatis,
30-90 x 3-4um. Mycelium secundarium superficiale: hyphis
cum conidiophoris primariis commixa in fasiculo per
stomatibus emergentis, pallide olivaceis, repentis, septat-
is, ramosis, subtiliter verruculosis, 2-3um latis,
conidiophoris secundaris lateraliter gerentis. Conidiis
cylindriaceis vel subcylindraceis, pallide brunneo-
olivaceis, minutissime verruculosis, rectis vel leniter
curvatis, plerumque 3-6 septatis (raro 1-2 septatis), non
constrictis, ad apicem rotundatis, inferne lenitissime
attenuatis, cellula basali in hilum semitruncatum et
atrobrunneum, 20-65 x 2.5 - 4yum. Habitat in foliis vivis
Bougainvilleae spectabilis (Nyctaginaceae). G. Lim, IV 1980,
No. SU 95 (Herb. LAM, YEN #10586), typus.
Leaf spot distinct, brown, orbicular or irregularly
orbicular, surrounded by a definite dark brown margin,
oy
|
LO_m
CED
Fig. A-E, Stenella bougainvilleae Yen & Lim. A, Fascicle of
conidiophores and formation of external hyphae; B,
Geniculate conidiophore; C, External hyphae and formation of
secondary conidiophores; D, Branched conidiophore; E,
Conidia.
98
scattered, 1-7mm in diameter, often confluent on the
terminal part of the leaf. Caespituli hypophyllous,
invisible even under the hand lens. Primary mycelium
internal: hyphae almost colorless, septate, branched, 1.5-
2.-5um wide. Stomata not developed. Primary conidiophores
hypophyllous, 2-8 in fascicles emerging through the strom-
ata, simple or occasionally branched, erect below, attenuate
and denticulate above, pale brown olivaceous and paler
towards the tip, 3-8 septate, 0-3 geniculate, apex attenuate
and decorate with small dark brown conidial cicatrices,
30-90 x 3-4um. Secondary mycelium external: hyphae arising
at the base of primary conidiophores and emerging through
the stomata, pale olivaceous, repent, septate, branched,
finely verruculose, 2-3\m wide, bearing laterally the
secondary conidiophores, which are simimar in respect to the
primary conidiophores. Conidia pale olivaceous, cylind-
dric or subcylindric, finely verruculose, straight or
slightly curved, generally 3-6 septate (rarely 1 or 2
septate), apex rounded, base attenuate-subtruncate and
decorated with dark-brown conidial cicatrice generally not
constrict, 20-65 x 2.5-4um.
On Bougainvillea spectabilis Willd. (Nyctaginaceae), in
Holland Road, Singapore, G. Lim, April 1980, No. SU 95
(Herb. LAM, Yen #10586).
Rao (1962) has described in India a Cercospora
bougainvilleae P. N. Rao, parasiting also on Bougainvillea
spectabilis; but it differs from ours by its hyaline
conidia. On the other hand, Sobers and Seymour (1969) have
described a Cercosporidium bougainvilleae (Munt.) Sob. &
Seym. which differs from our fungus in having very dense
fascicles and well developed stromata.
Cercoseptoria Cordiicola (Yen) Yen, comb. nov.
=Pseudocercospora cordiicola (Yen) Yen, Gard. Bull.
Singapore 33:173,.1980;
=Cercospora Cordiicola (Yen) Yen, Rev. de Mycol. 32:
182, 1967.
=Cercospora cordae Yen, Rev de Mycol. 29: 216, 1964.
non Cercospora’ ‘cordae -Chupp, Mongr. Univ. 2. Ra scer.s,
22%, 2457) 91934.
On Cordia cylindristachya (Boraginaceae), in Singapore.
Literature Cited
Rao, P. N. 1962. Some Cercospora species from Hyderabad,
India. indian Phytopath’ PY5=shi2.
sobers, E. K. and C. P. Seympour. 1969. ‘Proes Pla. St. whoec.
Soc. 81:398.
Acknowledgements
We are grateful to Don R. Reynolds, Natural History Museum,
Los Angeles, for reviewing the manuscript.
MYCOTAXON
VW Ode Lis NO. Ll. pp. 991.02 October-December 1982
PHYTOPHTHORA OPERCULATA SP. NOV., A NEW MARINE FUNGUS
K.G. PEGG and J.L. ALCORN
Plant Pathology Branch,
Department of Primary Industries,
Indoorooptlly, Queensland, 4068, Australta
In 1980 a species of Phytophthora resembling P. vestcula
Anastasiou §& Churchland was reported to be associated with
trunk cankers and decayed absorbing rootlets of declining
white mangrove trees (Avtcennta martna (Forsk.) Vierh.) in
the Gladstone area of central coastal Queensland (Pegg,
Gillespie and Forsberg, 1980). During a study to determine
the distribution of this fungus in Queensland mangrove
communities, another Phytophthora with a unique method of
zoospore release was recovered on several occasions from
decayed white mangrove leaves submerged in sea water
(Pegg and Forsberg, 1981). As this fungus has sporangial
characters which differ from those of other Phytophthora
species, it is described below as new.
Phytophthora operculata sp. nov. Figures 1-3
Hyphae hyalinae, ramosae, demum septatae, 8-10 um
diam. Sporangtophora non ramosa, sympodialia, torsiva,
6-7 um diam. Sporangta subcylindrica vel anguste
ellipsoidea, ad apicem late complanata, non caduca, 30-
175 x 25-75 um, obturamento septali in sporangium usque
ad 13 um protrudenti, ad apicem circumscissa. JZoosporae
flagellis binis lateralibus, limoniformes vel fusiformes,
globosae 10-12 um diam. ubi incystatae. Reproductio
sexualis ignota. Hab. in foliis putridis Avtcenntae
martnae, Moreton Bay, Queensland, 18.vi.1980, K.G. Pegg,
BRIP 13362, holotypus; IMI 249911, isotypus.
100
Fig. 1. Line drawings of Phytophthora operculata
sporangia. A. Mature sporangia before zoospore release.
B. Very young sporangium on spirally twisted sporang-
iophore. C. Sporangia after zoospore release (scale =
25 pum.
On solid media the mycelium is composed of freely
branching hyphae 8-10 um wide, non-septate when young
but developing septa with age. Sporangia are produced
Sparingly on solid media in the absence of free water,
and more abundantly when discs of V-8 juice agar or
naturally-infected host tissue are immersed in autoclaved
101
Fig. 2. Mature sporangium with differentiated zoospores,
and sympodially elongating twisted sporangiophore (scale
=e omni). Fig. 35. joporangium after dehiscence, with
prominent basal plug (scale = 25 um).
sea water. They are borne in a lax monochasial sympodial
arrangement on sporangiophores 6-7 um diam., and which
are in some parts twisted spirally (Fig. 2). Sporangia
are non-deciduous, mostly narrowly ellipsoid, sometimes
almost cylindrical, occasionally narrowed at the base,
flat and broad at the apex, smooth-walled, 30-175 x 25-
75 ym (Fig. 1). There is a conspicuous basal plug which
protrudes 2.5-13 4m into the sporangium. Zoospores
differentiate fully within the sporangium. A circumcissile
split occurs near the sporangium apex, allowing an
operculum to open and release the zoospores without
formation Of a vesicle. “The edges of the operculum
sometimes curl (Figs. 1, 3), but sporangia do not collapse
after dehiscence. Radial growth on corn meal agar after
5 days at 35°C was 4-7.5 mm, with no growth ate oe. Cee The
minimum temperature for growth was 18 C, and the optimum
in the range 21-31°C.
Sexual reproduction has not been observed. Oogonia
and antheridia were not produced when isolates were grown
in single culture, or when six isolates of the species
were paired in all combinations on 20% V-8 juice agar,
carrot agar, media containing B-sitosterol (Ribeiro, 1978),
or on detached white mangrove leaves incubated in auto-
claved sea water.
102
Isolates of Phytophthora operculata have been
obtained from decaying white mangrove leaves collected
from several localities in or adjacent to Moreton Bay,
south-eastern Queensland. It differs from all other
species described in the genus (Ho, 1981) by the
presence on the sporangium of an apical lid which opens
prionyto ‘zoospore release, . Other species uch) as
P. bahamensts Fell §& Master, P. eptstomium Fell & Master,
and P. mycoparastttca Fell & Master also have unique
methods of zoospore release (Fell and Master, 1975). It
seems that marine Phytophthora species are likely to
produce sporangia with characteristics not seen in
terrestrial members of the genus.
Cultures have been deposited as IMI 249911 and
ATCC 44952.
References
Fell, J.W. and Master, I.M. 1975. Phycomycetes
(Phytophthora spp. nov. and Pythtum sp. nov.) associated
with degrading mangrove (Rhtzophora mangle) leaves.
Canadian Journal of Botany 53: 2908-2922.
Ho, JH: 19Si° SSynoptic keys to othe speciessor
Phytophthora. Mycologia 73: 705-714.
Pegg, K.G. and Forsberg, L.I. 1981. Phytophthora in
Queensland mangroves. Wetlands 1: 2-3.
Pegs; K.G., Gillespic, N.C. ard Forsberg, sls ooe.
Phytophthora sp. associated with mangrove death in
central coastal Queensland. Australasian Plant
Pathology 9(3): 6-7.
Ribeiro, O.K. 1978:°.A source book of the’ genus
Phytophthora. J. Cramer, Vaduz.
MYCOTAXON
VOdee Ale Not lee pp... 05-106 October-December 1982
THECAPHORA ANDROSACINA AND ENTYLOMA
GAILLARDIANUM, NEW SPECIES OF USTILAGINALES
KALMAN VANKY
Tegelbruksvagen 1, 780 41 Gagnef, Sweden
The types of Thecaphora androsaces (Karsten) Gutner and
Entyloma gaillardtae (Speg.) Speg., proved not to be smut
fungi. Consequently, it was necessary that the smuts inha-
biting Androsace and Gatllardta be described as new.
THECAPHORA ANDROSACINA Vanky, sp. nov.
Typus: Androsace maxtma L., Hungaria, Comtt. Fejér, pr.
pag. Erd, "Kutyavdr", V.1874, leg. J. A. Tauscher (Herb.
Vertiag. Vanky no. 10751 -in UPS).
Sort multttudinem granulosam-pulveream sporarum conglo-
batarum formantes. Glomerult sporarum globost, ovotdet
usque parum elongatt vel forma trregulares, 16-85 x 20-45
(—52) pm e sports 2—-15(—25?) factle dtscedenttbus composttt.
Sporae rotundato-polygonales, trregulares, cunetformes vel
elongatae, 8-15 x 11-19 wn, flavidulo-hyalinae usque dtlute
flavtdulo-brunneae, partete 0.81 ym crasso, tn superftcte
contactus levt, tn parte extrorsa dense et trregulariter
verrucoso.
Sori in the seeds forming pale brown, granular-powdery
mass of spore balls. Spore balls (Fig. 1) globose, ovoid
to slightly elongated or irregular, 16-35 x 20-45(—52) um,
composed of 2-15(—25?) easily separating spores. Spores
(Fig. 1) polyangularly irregular with rounded edges,
cuneiform or elongated, 8-15 x 11—19 pm in diameter, yel-
lowish-hyaline to pale yellowish-brown, wall 0.8—1 um
thick, smooth on the contact surfaces, coarsely and irregu-
larly verrucose on the free surface.
Type on Androsace maxima L., Hungary, Distr. Fejér, near
the village Erd, "Kutyavar", V.1874, col. J. A. Tauscher
(Herb. Ustilag. Vanky no. 10751, located in UPS).
104
Several smut names have been mentioned in the literature
On Androsace, but they were either based on non fungal ma-
terial or nomenclaturally incorrect. Thus, Karsten (1907: 4)
described Usttlago androsaces in the ovaries of Androsace
2fitltformis Retz. as "Sori fusci, pulverulenti. Sporae
sphaeroideae, laeves, 35-50 yp diam.". Liro (1924: 343)
examining the original specimen of Karsten found that it
only contains young seeds which were mistaken by Karsten
for smut spores. Lavrov (1936: 31) found a smut forming
loose spore balls in the seeds of Androsace maxima L.,
considered it identical with Usttlago androsaces and made
the combination Sorosporium androsaces (Karsten) Lavrov.
Gutner (1941: 191) transferred this species to Thecaphora
as T. androsaces (Karsten) Gutner. Savulescu (1957: 869),
though aware of Liro's finding, used the illegitimate name
Thecaphora androsaces (Lavrov) Gutner. Another binomial for
an Androsace smut is the herbarium name "Theecaphora jubtlet
Jacz. n. sp. ined in herb. VIZR", mentioned by Lavrov (1936:
31), which in his opinion is identical with Sorosportum
androsaces.
ENTYLOMA GAILLARDIANUM Vanky, sp. nov.
Typus: Gatllardia aristata Pursh, (cult.) Romanta,
franssy wana, “Tirgu-Mures, QUp.i 52 0. te Ss Mle ie el ee
leg. K. Vdadnky. Holotypus: Herb. Usttlag. Vdnky no. 743
in UPS; tsotypt in Vdnky, Vet. 44 et tn Herb. mye. roman,
1725 (sub Entyloma composttarum).
Sort conspteut steut maculae dtspersae vel gregartae
foltorum diametro 0.1—5 mm vel confluentes majores, tnttto
dtlute flavtdulo-viritdes deinde brunnet, saepe cum margtne
tenut flavtdulo. Sporae globosae usque forma trregulares,
magnttudine variae, 9-15 pm tn diam., subhyalinae usque
flavae, partete levt, saepe bitstratoso, 1—2(-4) ym crasso.
Sori in the leaves as circular, scattered or gregarious
spots of 0.1—5 mm in diameter, or more by confluence, at
first pale yellowish-green, later brown, often with a thin
yellowish margin. Spores (Fig. 2) globose to irregular,
variable in size, 915 um in diameter, subhyaline to yellow,
with smooth, often two-layered, 1—2(—4) ym thick wall.
Spegazzini (1925: 148) described Entyloma gatillardtae
(Speg.) Speg., based on Protomyces? gatllardiae Spegazzini
(1909: 284) on Gatllardta donitana, and characterized by
spores 30-35 pm in diameter. Spegazzini (1909: 284) himself
105
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~-
oO WV
tek
oP.)
Qa
N~—
ewe,
Fig
LM
(type) in
tanum
Llard
10 pm.
Fig. 2. Spores of Entyloma gat
Bar
106
suspected that this fungus could represent oospores of
Peronosporales. This opinion was also supported by Savile
(1947: 117), Zundel (1953: 251) and Lindeberg (1959: 36).
The true Entyloma smut on Gatllardta spp. was therefore
referred by most of the authors to the collective species
E. composttarum Farlow. However, by comparing Entyloma
on Gatllardia aristata with EF. composttarum (on Aster
puntceus L., USA, Massachusetts, Wood's Holl, IX.1883, W.
Trelease, in Ellis, N. Amer. fgi. 1492), I found that the
first species has more irregular and somewhat larger
spores, with thicker and often two-layered wall. In my
Opinion these are sufficiently distinct to consider the
Entyloma on Gatllardta aS a separate species.
ACKNOWLEDGEMENTS
The author would like to express his sincere thanks to
Prof. J. A. Nannfeldt and Dr 0. Constantinescu (Institute
of Systematic Botany, Uppsala, Sweden) for the reading of
the manuscript, and to Dr S. T6th (University of Agri-
cultural Sciences, Gddd116, Hungary) for the preparation
of the Latin description.
LITERATURE CITED
Gutner, L. S. (1941). Golovnevye griby. Moscow, Leningrad.
Karsten, P. A. (1904). Fungi novi, paucis exceptis, in
Sibiria a clarissimo 0. A. F. Loennbohm collecti.
Oevers. Forh. Finska Vetensk.- Soc. 46(11): 1-9.
Lavrov, N. N. (1936). Novye i bolee redkie golovnevye griby
sem. Ustilaginaceae severnoi tsentralnoi Azii. Trudy
Biol. Nauchno-Iss]. Inst. Tomsk. Gosud. Univ. 2: 1-35.
Lindeberg, B. (1959). Ustilaginales of Sweden. Symb. Bot.
Upsa ln 16(2) Sl 17.5%
Liro, J. I. (1924). Die Ustilagineen Finnlands, I. Ann.
Acad. (S¢1.) Fenn, Ser. A, 1621-036.
Savile, D. B. 0. (1947). A sudy of the species of Entyloma
on North American Composites. Canad. J. Res. Sect. C,
BOs Scie con= 105-120:
Savulescu, T. (1957). Ustilaginalele din Rep. Pop. Romina.
Vol. 2. Bucuresti.
Spegazzini, C. (1909). Mycetes argentinenses. Anales Mus.
Nac. Hist. Nat. Buenos Aires, Ser. 3, 12: 257-458.
(1925) Ustilagineas Argentinas nuevas o criticas.
Revista Argent. Bot. 1: 145-156.
Zundel, G. L. (1953). The Ustilaginales of the World.
Contr.Dept. ‘Bot..Penns. State. Collie No.ieGc e410:
MYCOTAXON
VO v to NOS Lp. Di pads October-December 1982
A NEW SPECIES OF ORBILIA FROM CANADA
J. We HAINES.
Btologtecal Survey, New York State Museum
The State Educatton Department
Albany, New York 12230
K. N. EGGER@
Btosystematics Research Instttute, C.E.F.
Ottawa, Ontarto
Canada KlA 0C6
ABSTRACT
Orbilia piloboloides, a new species of Helotiales
from the bark of Ulmus americana, is distinguished by
its subapically swollen paraphyses which resemble the
sporangiophores of Pilobolus. An anamorph with what
appear to be sympodially produced blastoconidia is pro-
duced in culture.
Orbilta is one of the most easily recognized genera
of Helotiales, yet relatively few treatments of the genus
have been published (Dennis 1978, Svréek 1954, Seaver
1951) and many of its species are poorly known. Despite
this, one taxon, represented by some recent Canadian col-
lections is so markedly distinctive in having subapically
swollen paraphyses and nearly fusoid spores that it is
described here as a new species. All three collections
are from relatively undecayed bark of Ulmus amertcanus L.
and not on well-decayed, decorticated wood or herbaceous
stems as is usual for Orbiltia.
Dm bidahed as New York State Museum Journal Sertes #347.
{Present address: Department of Biology, University of Victoria,
P. O. Box 1700, Vietorta, BC, Canada V8W 2Y2.
108
The fungus was obtained in pure culture by placing a
moistened apothecium on the lid of an inverted petri dish
and allowing the ascospores to shoot upward onto the sur-
face of 1.25% Malt agar. A characteristic anamorph was
observed in culture.
ORBILIA PILOBOLOIDES Haines and Egger, spec, nov,
Apothectts sesstltbus ad brevi-stipttatis, qurantt-
lutets. Ascts (35-) 40-52 (-55) x 3-4 ym, Ascosports
8-10 x 2,0-2.6 ym,, fustformtbus ad navtcultformtbus.
Paraphystbus filtformtbus cum bulbo aptcalt crassttuntcato
et cum gtbba subapticalt pyrtformi tenuttuntecato, 2,0-3,3
wn. diam.
Colonits lentis crescenttbus, hyqlints ad aurantts,
rugulosts, semimucosts. Mycelits hyphotdets vel monilt-
formtbus, ramosts, laevtbus, hyalints. Contdtophorts
sympodtaltbus, 5-12 (-20) x 1,5-3 wm, Sympodulocontdits
laevtbus, hyalints, curvts, l-septatts, 9-12 x 2-3 um,
Holotype: DAOM 176753. Ginns' Farm, St. Eltzabeth Road,
Cantley, GattnequCo., Quebee, Canada, April 22, 1980,
J. H, Ginns, (100+ apotheecta on Ll ptece of substrate,
lL dried culture of anamorph, living culture tn DAOM
culture collectton).
TELEOMORPH: Apothecia (Fig. 1) scattered to gregarious on
relatively undecayed bark, luteus when hydrated, darkening
to orange-luteus on drying, waxey-appearing, often with a
glaucus coating most noticeable as a white fringe at the
margin, cupulate at first, becoming planate at maturity,
up to. 1.5 mm diam., folding or curling inwards, at the mar
gin on drying, sessile to short-stipitate. Stipe, when
present, 0.2-0.3 mm diam, by up, to 0.3 mm high, Pignter—
colored than cup, sometimes with a restricted web of hyphae
at the base. Exciple (Figs. 6 and 8) composed of thin-
walled, hyaline cells 5-15 um across forming textura angu-
laris. Asci (Fig. 3) (33-) 40-52 (-56) x 3-4 um, 8-spored,
cylindrical in upper two thirds, tapered in lower third to
a contorted base 1-2 um across, apparently not subtended
by croziers, distinctly truncated and sometimes slightly
enlarged at the apex, without visible pore or amyloid re-
action, thin-walled throughout. Spores (Fig. 4) 8-10 x
2,.0-2.5 um, short-naviculate to fusoid, bilaterally sym-
metrical, slightly more acute at the apex than base, non-
septate, smooth, thin-walled, without conspicuous internal
Figs. 1-8 Orbilia piloboloides, 1, Apothecia on natural substrate,
approx, 12X, 2, Hymenium 500X, 3. Asci 1,000X. 4. Ascospores 1,000X,
5. Paraphyses 1,000X. 6. Section of apothecium 400X. 7. Asci and
paraphyses 1,000X. 8. Detail of ectal exciple 1,000X, (A111 from the
holotype).
TELUS)
110
features. Paraphyses (Fig. 5) filiform, 0.9-1,2 wm diam.
in the lower portion, swollen at the apex to a distinct
spherical, thick-walled knob 1,1-1.8 um diam. subtended
by a thin-walled, pyriform swelling 2.0-3,.3 um diam.,
often with a few amorphous particles adhering to the sur-
face, extending beyond the asci by a few microns in the
hymenium. Hymenial components not adhering, easily separ-
ated by pressure on squash mounts (Fig. 2, 7).
ANAMORPH: Colonies submerged or appressed to the surface
of agar, rugulose, dense, surface somewhat slimy, hyaline
on 1.25% malt agar and potato carrot agar to white on po-
tato dextrose agar, developing a light orange pigment when
exposed to ultraviolet; slow-growing (6-10 mm in 10 days
at room temperature). Hyphae (Fig. 9) hyaline, smooth,
sparingly septate, branched, 1.2-2.5 um wide, occasionally
interspersed with moniliform mycelium with elliptical
swellings up to 4 um wide, septate at 4-8 um intervals and
constricted at the septa, Conidiophores (Figs, 710,71), 90)
sympodial, hyaline, lageniform to cylindrical, 5-12 (-20)
x 1.5-3 um, arising from undifferentiated mycelium, usually
with one to three denticles at the apex or occasionally
with an elongated rachiform apex. Sympoduloconidia (Fig.
13) apparently blastic, smooth, hyaline, strongly curved
with an obtusely rounded apex and a narrowly truncate base,
9-12 x 2-3 um, with a single median septum, not constricted.
SUBSTRATE AND HABITAT: Bark of mature, standing or disin-
tegrating Ulmus amertcana killed by dutch elm disease. Often,
if not always, originating from the upper parts of the
trees (J. Ginns, personal communication).
ETYMOLOGY; piloboloides = Pilobolus -like. Refers to the
paraphyses which resemble the sporangiophores of the zygo-
mycete Ptlobolus Tode,.
COLLECTIONS EXAMINED: CANADA: Quebec: Ginns' Farm, 45°
34'N, 75°47'W, St, Elizabeth Road, Cantley, Gatineau Co,
on bark of log of Ulmus amertcana 22 April 1980, J. H. Ginns,
DAOM #176753 (Type) DAOM, (Isotype) NYS 26 October 1980,
J.-H. Ginns DAOM #177611, (DAOM, NYS). Ontario 3 north
Gower, on inner bark of log of Ulmus americana, 16 July 1979,
G. P. White #169, DAOM #176754 (DAOM).
DISCUSSION: This species is immediately recognizable as a
member of the Orbiliaceae by its waxey, orange apothecia,
its small flat-topped asci without apparent pores, knob-
tipped paraphyses and globose or angular-celled excipulum.
Figs, 9-13 Orbilia piloboloides, 9, Germinating ascospores, 10, 12,
Sympoduloconidia and conidiophores of anamorph, 11. Conidiophore
apices showing rachiform development, 13, Mature conidia, A11 at
1,000X, 9, from DAOM 17764, 10-13. from culture of holotype,
plate
~
LZ
It is distinguished from all other species by its unusual
paraphyses which resemble sporangiophores of the zygomycete
Pilobolus. The ‘similarity is entirely superficial, however,
and there is no evidence that the paraphyses are directly
involved in dispersal. The ascospores are small and non-
septate as in other orbilias but they have a shape not
previously described for any other member of the genus,
Recent studies (Benny, Samuelson and Kimbrough 1978)
have demonstrated the presence of a blue-green algal (Cyan-
obacterial) associate in the excipular tissue of Orbilta
luteorubella (Nyl.) Karst. Although one of us (JH) has con-
firmed these findings and has observed algal cells in
several other species of Orptlta, none were found in 0.
ptlobolotdes. The presence of algal cells in the lower apo-
thecial tissues of orbilias appears to be correlated with
conditions suitable for an abundance of photosynthetic
organisms on the substrate surface, Orbilia ptloboloides
occurs on substrate relatively poor in these organisms,
Hymenial components which are firmly held together in
a glutinous matrix are common for some species of Orbilta
but those of 0. ptlobolotdes do not appear to be embedded
and are easily separated by applying pressure on squash
mounts, Stipitate species are rare in the genus, Orbtlta
ptlobolotdes often has a short stipe, but it is a variable
character which may be influenced by the environment near
the substrate,
We do not feel that the anamorph of 0, ptlobolotdes can
be accommodated in any known genus, However, there does
not .seem to be acase at this time for, ecrecuing fa anew
genus,
Berthet (1964) described an anamorph of Orbilta xantho-
stigma (Fr.) Fr. with conidia of two types: fusoid didy-
moconidia and "horseshoe-shaped" bilobed conidia, This
anamorph has been referred to Dtcrantdton Harkn, (fide Henne-
bert and Bellemere, 1979). Butterfield (1973) also noted
the presence of lobed conidia and phragmoconidia in D,
fragtle Harkn, The anamorph of 0, pilobolotdes does not have
bilobed conidia and the conidia are curved rather than
fusoid, although the method of conidium production is simi-
lar to Dteranidton, Dtplorhtnotrichun Héhnel (= Dactylarta Sacc,,
fide Bhatt and Kendrick, 1968) has been used for a group of
fungi with hyaline didymoconidia on sympodial conidiophores,
but the conidia are not curved, Idriellqa Nelson and Wil-
helm, which has a Hymenoscyphus teleomorph (Kimbrough and
Atkinson, 1972), and Miecrodochtum Syd, (Sutton, Pirozynski
and Deighton, 1973) are similar fungi. JIdrtella spp, produce
dark chlamydospores in culture and has a faster growth rate
than our species, (Orbiltq ptlobolotdes produces moniliform
hyphae in culture but these do not become pigmented,
Microdechtum is known primarily from the natural substrate,
At least one species, Mierodochtum phyllantht Sutton, Pirozynski
and Deighton, does not produce chlamydospores in culture
Las
but does produce a dark brown pseudoparenchymatous stroma,
Orbtlia ptlobolotdes produced no stromata in culture,
ACKNOWLEDGEMENTS: The authors wish to thank Dr, J. D.
Bissett for his advice and discussions during the develop-
ment of this paper and Dr. J. Ginns for kindly offering
fresh collections for culturing, Thanks also to Drs, Scott
Redhead and James Kimbrough for their helpful comments,
BIBLIOGRAPHY
Benny, G. L., D. A. Samuelson and J. W. Kimbrough. 1978, Ultra-
structural studies on Orbtlta luteorubella (Discomycetes). Can.
Smeoc, 56:2006-2012,
Berthet, P. 1964, Formes conidiennes de divers discomycetes. Bull,
SOG. Mycol. £r. O03126.
Bhatt, G. C. and W. Bryce Kendrick, 1968. The generic concepts of
Dtplorhtnotrichum and Dactylarta, Can. J. Bot. 46:1253-1257,
Butterfield, W. 1973. Morphological variation of Dteranidton fragile
and D, tnaequalis in culture. Can, J. Bot, 51:795-799,
Dennis, R. W: G. 1978. British Ascomycetes. Cramer, Lehre,
Hennebert, G. L. and A. Bellemere. 1979. Les formes conidiennes des
discomycetes. Rev. Mycol. 43:264,
Kimbrough, J. W. and M. Atkinson. 1972. Cultural features and imper-
fect stage of Hymenoscyphus caudatus. Amer, J. Bot. 59:165-171,.
Seaver, F. J. 1951. The North American Cup-fungi (Inoperculates).
Seaver, New York.
SULEONs be Cs, Ky As ePirozyuski and fF, o Deighton. 1972, Mtcro-
dochtum. Syd. Can. J. Bot. 50;1899-1907.
Syrtek, M. 1951, Rivyise Velenovského druht rodu Orbilta (Discomy-
cetes). Rivisio critica J. Velenovskyi speciarum generis Orbtlta.
Sb. Narod. Mus. Praze 10B:1-25,
MYCOTAXON
VOL WAVE p NOt pe bao LG October-December 1982
PHELLINUS ANDINA PLANK & RYV. NOVA SP.
by
Sie Plank! Ayal diye Ryvarden2
ie InStireut, £. Umweltwissenschatten uu. Nacubnschuez
der) Oster. Akad.’ Wiss.,, H@INrILChsStr4 5/7 bat
A-8010 Graz, Austria.
2. Botanical institute, University obe0slo,
P.O. Box 1045, Blindern,. Osilo.s, Norway.
Abstract
Phellinus andina is described from Argentina. It is
characterized by a resupinate fruitbody, short tramal setae
in the dissepiments and large, subqlobose, golden brown
spores.
Under an expedition to Argentina, one of us (S. Plank)
collected a Phellinus species with a unique combination of
characters and for which we could not find a name.
Phellinus andina Plank & Ryv. nova sp.
Fructificatio resuninata, effusa, adnata marginibus sub-
tiliter .floccosis, 1-3.-mm latis, vallide® tabacinis. Facies
pororum umbrina poris rotundis,~-6-7 per mm, tubis concolo-
ribus, ad 500 um altis. Subiculum cinnabarinum ad tabaci-
num. - Systema hypharum dimiticum: hyphae sceletales crasse
tunicatae, aureae, alterae in trama, alterae in tubarum
dissepimentis sitae, 2-3.5 um latae, hyphae in subiculo et
in margine sitae ad 5 um latae luminibus latioribus. Hyphae
generatoriae hyalinae, septis simpliciter septatae, 2-3 um
latae. Setae vel hyphae setoideae in dissepimentis adsunt,
breviter emergentes, pullae, crasse tunicatae, rectae vel
leviter curvulae, 4-7 um latae, 35-60 um longae. Setae
hymeniales desunt. Basidiae claviformes, 10-20 um longae,
4-6 um latae, sterigmatibus 4. Basidiola evoluta hyalina,
DV |
Pro. io, .Pnetlinus; andina. A) Section, «through? the. disse-
piments, B) skeletal hyphae from the trama, C) setae from
the dissepiments, D) basidiols, E) basidium, F) spores.
From the type.
L16
10-20 um longa, ventricosa. Sporae subglobosae, ad late
ellipsoideae, crasse tunicatae, aurei-umbrinae, 7-9/5.5-7
um.
Typus: Ad truncum mortianum Myrceugenellae apniculatae
(Myrtaceae). In Argentina, regio Rio Negro (Andes),
"Bosque de Arrayanes" apud lacum "Nahuel Huapi",
ca. 800 m. Leg. S: PLANK & M. FP. BROGGT, 26.01. boa
HOw iy Omiya) Wr Ses GZUe
I seo.t yop 1: O8lo; Ky BPI, “Coll. aucte Gra e-aAl A RG
Ds
Fruitbody resupinate, effused and adnate. Margin finely
floccose, 1-3 mm wide, pale snuff brown, pore surface umber
brown, pores round, 6-7 per mm, tubes concolorous, —Up te
500 um deep, context deep cinnamon to snuff brown, dense
and up? to I smm thick... Hyphal’ systems dimitic, Sskeletar
hyphae thick-walled and golden brown, in the trama 2-3.5 um
wide; in the context and margin up to 5 um wide: and iwwithua
wider lumen. Generative hyphae hyaline, simple septate and
2-3 um wide. Tramal setae or short setal hyphae present in
the dissepiments, slightly projecting, dark brown, thick-
-walled, straight to somewhat sinuous 4-7 um wide, 35-60 um
long. Hymenial setae absent, Basidia clavate 15-20 x 4-6 um
with 4 sterigmata, basidiols present, ventricose, hyaline
10-20 um long, spores subglobose to broadly ellipsoid,
thick-walled and golden brown by maturity, 7-9 x 5.5-7 um.
Only known from a died stem of Myrceugenella apiculata
(Myrtaceae).
The short tramal setae restricted to the disseniments make
this species unique in Phellinus. Such setae are known in
Inonotus, such as in I. hastifer Pouz.; but thave nithepta
been unknown in Phellinus. Furthermore, the large spores
are also unusual in the genus where most species have
spores shorter than 7 um. The new species reminds in seve-
ral respects about an Inonotus species, but the distinct
dimitic hyphal system with thick-walled, narrow skeletal
hyphae rules out a place in the genus.
Acknowledgements.
The first author (P) wishes to express his. sincere
thanks to Mario: F. Broggi, Vaduz, for valuable assistance
during the excursion to Argentina and to the Amt der Steier-
markischen Landesregierung, Abteilung ftir Wissenschaft und
Forschung £or. financial ‘support.
MYCOTAXON
Vouyer Vs (Notas opp eal 7122 October-December 1982
A REINVESTIGATION OF THE NORTH-WEST HIMALAYAN PULVINULAS
RISHI KAUSHAL
Department of Botany, Panjab University, Chandigarh 160014, India
SUMMARY
On a reinvestigation, it has been found that the N.W.
Himalayan specimens of Pulvinula Boud. belong to P. mussoor-
tensts (Thind, Cash §& Singh) Batra, P. globtfera (Berk. §&
Curt.) Le Gal, P. ortichaleea (Cke.) Rifai, P. convexella
(Karst.) Pfister, P. laeterubra (Rehm) Pfister, P. nepalen-
sts R. Kaushal sp. nov., Pulvinula taxonomic sp. I and
Pulvtnula taxonomic sp. II. A key to all these species is
given.
Species of Pulvinula Boud. from N.W. Himalayas have
been poorly understood. For example, Thind & Batra (1957),
Thing q Singh (1959) and Thind et al. (1959) probably were
not clear about its generic concept when they treated species
Of Pulvinula under Lamprospora de Not. _ Later, Batra q@ Batra
(1963) evidently were confused over these two genera. They
ducted. cOmeuspecies under Puivinula butyat the same time
deSeripnediat east one sor its Species Under Lamurospora.
Subsequently, more Pulvinula species were described from the
N.W. Himalayas by Thind §& Waraitch (1970) and Waraitch §
Thind (1977). ‘The tecent delimitation of species of this
eentisopy Ritar (1968), Moravec (1969)" and Pfister (1972,
Ie7oji end the importance of Critical mrcroscopic observa-
Ctonsy or proper identification led the author to re-examine
collections from the N.W. Himalayan ranges of India and
Nepal where the earlier Indian researchers made their
collections. An attempt has been made here to place the
collections (both published and unpublished) in the species
as understood now.
ASy a eresult ior these investivations, 1e has been fLound
that from among the earlier reports of the Pulvinula species
from N.W. Himalayas, only P. mussoortensis (Thind, Cash §
Singh) Batra was valid. Besides P. mussooriensis, N.W.
Himalayan specimens have been found to represent seven more
SeccClcs VizZ., £5 gLloortera tBerk. G iCurt..) Le, Gal, &
evtcuaueea (CKke.) Rita, PP. "converel la (Karst.) Prister,
Pa bacterupra (Rehm) Pilister, ©. nepalensts KR. Kaushal, sp.
NOV... Lulvinutla taxonomic sp. I and Pulvinuto taxonomic sp.
LES
A key to all the N.W. Himalayan species of Pulvinula
has been provided in this work. Comments have been included
for the collections which are at minor variance with the
118
described species. Collections have been commented under
the species to which they were assigned by the earlier
workers.
Pulvtnula cearbonarta (Fuck.) Boud. var. brevtspora Batra
Thind §& Batra (1957) first described this fungus (No.
154) as Lamprospora ecarbonaria, but later Batra in Batra §&
Batra (1963) shifted it to Pulvinula and raised a new variety
brevtspora. The collection is in decayed (condition an
preservative and, except for the size of ascospores (up to
14 wm across) and filiform, curved paraphyses which are not
enlarged apically, nothing more could be made out of it.
Going by Thind §& Batra's description, except for the pyvo-
philous habitat it does not seem to be different from P.
ortechalcea (Cke.) Rifai. Specimen examined: India: Batra
154 (PAN), on charcoal beds which were overgrown by mosses,
Jabber: Khet....Mussoorac, August 20,4. i95Z.:
Pulvinula constellatto Boud.
This species has been tentatively regarded as a synonym
of P. convexella (Karst,) Pfister by “Pfister (1970)2 sGUe.
In 27 referred to Py “constellatio by Batrarg Batraytigos)
has ascospores 15.8-18 um across, asci 194-248 um long, with
regular croziers and filiform, curved and branched paraphyses.
In my opinion 2t represents P&.. converetia. specimen
examined: “India: Batra’’27-(CUP-In) ,(on ‘soni. Dhobe Guaw
Mussoorrey UyP i, pen cemper LIo4.
Pulvinuta diseotdea (P., Henn. § Nym.) Batra
I follow Pfister (1976) who does not recognize this
species in. the absence, of its. type material. CUP-In 49
described by Batra §& Batra (1963) as P. dtscotdea has few,
small (about 1.5 mm in diam.) whitish apothecia, which are
not well preserved..)The asci are) 215-Z252ex%, 18-20 ime
spored, tapering below, withsregular croziers > suhesasco.
spores are 16-18 um across and have many small oil droplets.
The paraphyses are unbranched, up to 1.8 um wide at the base
and -are-up to 4. (=5) um wide above at their bent apices. main
ascospore size it is unlike any of the other white species
of this genus and it seems to be a new species (Pulvinula
taxonomic. sp. J) «sOQwing to Batra's insuhticient and poarm.
preserved material) 1 am not naming this species, Ihave
some good material of this fungus from the Eastern Himalayas
which would be published shortly. Specimen examined: India:
Batra 19 (CUP=In)); on soil, mossy falls, Mussoorie? Ucre
Pulvinula ertotara (Cke.) Le. Gal
The specimen used by Le Gal (1953) to describe P.
ettolata has. been referred to P. tetrasporas(Hanst shiver
by Rifai (1968) who has described its position in detail.
No. 2039 assigned to P. ettolata by Thind §& Waraitch (1970)
has small (up to 2.5 mm in diameter) whitish apothecia.
These bear 8-spored asci up to 235 wm long, taperingaeateene
base and arising from regularly two-pronged croziers, asco-
spores up to 12.8 (-13) um in diameter, paraphyses filiform
with curved apices up to 1.6 um wide throughout their length.
This Himalayan collection seems not to be different from
CL
Pogvoptyera (Berk. “G Curt.) Le Gal sensu Rifai, Though its
relatively smaller apothecia and ascospores, and the form of
the paraphyses bring it nearer to P. lacteoalba J. Moravec,
this species except for its 4-spored asci, may not be
different from P. globifera as noted by Pfister (1976).
Specimen examined: India: Waraitch 2039 (PAN), on humi-
colous soil amongst needles of some conifers under coni-
Forous forest, Baphr, Simla, HVP., August’ 19%) 1966.
Pulvinula globtfera (Berk. & Curt.) Le Gal sensu Rifai
PAN 2322 referred to this species by Waraitch § Thind
(1977) has yellow apothecia up to 3 mm in diameter and
Dy ropnilous. habitat... The asci are 180-218.x 14-18 um, 3-
spored, instead of tapering below become slightly narrower,
then abruptly contracted into a short cylindrical stalk-like
base agamiack regular croziers.. The ascospores are 12.8-
15.5 um across and the paraphyses are up to 1.6 um wide
DeLOow Pexpandingaup to Ss wm at’ their bent ‘to ‘curved. apices:
Its substrate, the type of ascal bases and size of
ascospores are unlike P. globtfera. Pulvinula neotroptca
Pfister, a pyrophilous species, is close to it but has pale
yellowish-greenish apothecia and smaller asci arising from
prominent croziers. The similar ascal bases, somewhat
Similar ascospore size and apically inflated paraphyses
alienate this Nepalese collection to some extent with P.
mereena (berk.,) Rifai, but the latter is distinct in* its
crimson apothecia, terricolous habitat and longer asci (up
LOmcGorum, 11de Ritai, 01908). (Waraitch’s collection ‘seems
to represent a new species for which I propose Pulvinula
nepalensis R. Kaushal sp. nov.* Specimen examined:
Nepal-ee Holotype: Waraitch 2322 (PAN), .on. .charcoal ‘and
burnt soil around a fire place in Bamboo grove, Sundarijal,
Kathmandu, August 15, 1969.
Pulvinula haemasttgma (Hedw. ex Fr.) Boud.
I follow Pfister (1976) and consider P. haemasttgma a
"nomen confusum.' Batra §& Batra recorded this fungus in
their check-list of "Indian Discomycetes" as P. haemato-
Prromamsich. Under thys species they. have moticited the
number of their collection deposited at PAN and it could not
be located there.
There are four unpublished collections at PAN, labelled
acm anaemasteomda. NO» 2252 and 2264 except: for their
smaller apothecia (up to 2 mm in diameter) and somewhat
smaller asci (204-240 x 17-19 um) are typical of P. con-
vexella (Karst.) Pfister in their basally distinctly forked
asci with regular croziers, size of ascospores (16.4-18.8
um across) and apically regularly branched filiform para-
physes.
*Apothecia ad 3 mm diam., disciformia, flavida. Asci 180-
205 x ATS sm, octosporz. Ascosporr 12..8=15.5) um diam,
globosi, laevigati. Paraphyses filiformae, ad 3 um ad
apices. Holotypus: Waraitch 2322 (PAN).
120
In No. 2470 “and 2562 ascospores are 14-18. 6, umtacrocs
and ascii, 2245-250. x 14.5-19 um which alsofallenineihe wage
of P. convexella, but their ascal bases instead of arising
from two-pronged regular croziers are simple, tapering and
lacking regular “croziers and :also. the regular Dranchanvaog
the. pataphyses as lacking in them. . These cotldenort abc
asSsSipnedetor Pe.) miliina because: of the tatters abruptety.
of ascal base and different color. PP. ortchalcea scensece
be the closest but has smaller ascospores. Probably these
two collections represent a new species (Pulvinula taxonomic
sp. ll). Specimens examined: India: Waraitch 2251) (Pane
on much’ wet, "soil ‘Harwan, Srinavar, J 1G Kj August, 7277 one
Waraitch 2264 (PAN), on much wet soil amid mosses along a
streamlet, Joru Pury, Biybihara-Pahaloam Road, a7.
September 24, 1967," 5. Chander 2470 (PAN) = yon wet worm,
Banikhet,” Dalhousie, HVP., August Tl, 19725 Ss. Chander
25627 (PAN) ,) on wet «soil, Kilbury, Nainital, UsP.. Aucus oe
BO.
Pulvinula haemasttgma (Hedw. ex Fr.) Boud. var. gtgantea
(Thind §& Singh) Waraitch § Thind
This variety, earlier published as Lamprospora haemato-
svigmaevar. gigantea by Thind "G Singh (1959) "to acconmmoda7:
No. 296, was later transferred to Pulvinula and validated
by Waraitch § Thind (1977) who also added two more collect-
DONS, ENO.) ZooemanGdeNOy 62955) CO) ete
In No. 296 the type of asci and paraphyses and! theisize
of ascospores are similar to those in P. ortchatcea and
should be taken as a representative of this species. No.
23385) 2559 alonetwith, Nos? 213i fand 2542 “(unpub lrchedmanc
deposited at PAN as. FP. haemastigma var. gigantea) in my
opinion also belong to P. ortchalcea. Specimens examined:
India: Singh 296 (PAN), on humus amid mosses in oak
forest, Brewery Road. Mussoorie, UlP. , AugustelU ooo.
Waraitch 2131 (PAN); on moist and sandy soil in mixed
forest. Pan} Pulla sDetnousie, (HYP. August) foo.
Nepal: Waraitch 2338 (PAN), oniwet clayey soil in angio-
spermous forest, Godaveri, Kathmandu, September, 170.
Waraitch 2339 (PAN), on wet sandy soil amid mosses in oak
forest, Daman, September 12, 1969; Waraitch 2342 (PAN), on
wet sandy soil in Oak forest, Daman, September 12, 1969.
Pulvinula mussooritensts (Thind et al.) Batra
This species, proposed by Thind et a1, (1969), has been
commented upon by Pfister (1976) who considered it to be
very near 2. niveocalba J... Moravec, “and I also concur wie
him. Specimen examined: India: Type: Singh 306 /(PAN))
on soil amid mosses, Brewery Road, Mussoorie, U.P., August
2 OSG
Pulvinula taxonomic sp.
Waraitch §& Thind.(1977)published this fungus togzccon-
modate Nos. 2327 and 2349. These have yellowish apothecia,
11.2-14 um across ascospores and up to 250 um long asci with
tapering base arising from distinct crozier., These appear
to represent the yellow form of P.. Laeterubra. | Specimens
examined: Nepal: Waraitch 2327 (PAN), on wet clayey soil
Del
in angiospermous forest, Balaju, Kathmandu, August 22, 1969;
Waraitch 2349 (PAN), on wet clayey soil in angiospermous
POrest., Balaju,Kathmandu, september 25, 1969.
Lamprospora multtguttulata Batra
CUP-In 104, the holotype of L. multtguttulata, which I
received through the kindness of Prof. Richard P. Korf con-
Sists of only a slide of a vertical ‘section of an-apotheciun.
However, the sections are poor, and the ectal excipulum
described as being compact textura intricata by Batra (1960)
could not be confirmed. The ascospores are smooth, multi-
guttulate and the paraphyses are slightly. bent fully
agree with Rifai's annotation slip in the herbarium packet
that CUP-In 104 is in no way related to Lamprospora and
instead represents some species of Pulvinula. Since its
microscopic features could not be fully reinvestigated, no
attempt is made to place it. Specimen examined: India:
PVOG weoatra Lod (CUP—In), “onssorl. Mussoorie, UP.
KEY TO THE N.W. HIMALAYAN SPECIES
Apothecia white ee hae ea oe
Apothecia yellow, orange or reddish
a)
W bo
yeeascospores up to 12.8 (-13) um in diameter; para
physes up to 1.6 ym wide Cpa eid their
Pength” —i 3 te at ee CCl tera
2. Ascospores 16- 18 um in ‘diameter; “paraphyses
up to 1.8 ym wide at the base and up to 4
(-5) wm wide at the apex .. Pulvinula taxonomic sp. I
J eASCL Without prominent, two-pronged ‘croziers . . «-. . 4
Stoel whl OTOmMImMenten CWO: Dron gedNcroziers:) S093 (be i186
Apothecia yellow; asci’up to 218 ym long,
abruptly contracted below into short, stalk-
ike base! .» . + ERC ee eg eats) LeU a@Lenetis
4, Apothecia salmon to orange: aS Ci. up to 260
EN, SONA CEeE pee hove et Bey cee memes 5: 0d ote bck < Carmen ee Sa ie ec eae Pe Pea)
SaerescOspores up tO; LS fim in “diameter 2 20 2. corveralcea
Syenocospores. 14-186 “mein
Gtanecet. uu, os -. Peete. 6 EE aenw La CaxOnoOlLe Sp. (itl
6. Ascospores 16.4-18.8 ym in diameter;
paraphyses regularly branched at their
AUEOOS ead ese em hy Noa Ame ed te eth are fet KOON DONC T EG
6. Ascospores smaller; paraphyses branched
OceUN De AICIG (29 "ha, Moree Vea. Gas Sk te ee RN Te Seite l ae Gy 27.
7. mscospores 11-14 ym in diameter; paraphyses
often branched .., : P. laeterubra
7. Ascospores 9-11.5 um in diameter: paraphyses
WTC ed AAS nee i a spears . P. mussoortensts
ACKNOWLEDGEMENTS
I am deeply indebted to Prof. Donald H. Pfister, Farlow Herbarium,
U.S.A. for carefully checking this manuscript, loaning some specimens of
P22
Pulvinula, and for all his valued suggestions leading to the completion
of this manuscript. I wish to thank Prof. Richard P. Korf for the loan
of specimens from CUP. To Prof. K. S. Thind, my revered teacher, I am
very grateful for constant encouragement and for his permission to study
the collections at PAN. The research was financed by Department of
Science and Technology, Government of India.
REFERENCES
Batra, L. R. 1960. New species of Discomycetes from India. Il.
Mycologia 52:524-527.
and S. W. T. Batra. 1963. Indian Discomycetes. Univ. Kansas
Sci. Bull. 44:109-256.
Le Gal, M. 1953. Les discomycétes de Madagascar. Prodr. Flore Myc.
Madagascar 4:1-465.
Moravec, J. 1969. Nekteré operculdtni diskomycety nalezené v
okresech Mlad4 Boleslav a Jicin. Cesk& Myk. 23:222-235.
Pfister, D. H. 1972. Notes on Caribbean Discomycetes-II. Two species
of Pulvinula from Puerto Rico. Phytologia 24:211-215.
1976. A synopsis of the genus Pulvinula. Occasional Papers ‘of
the Farlow Herb. of Harvard Univ. 9:1-19.
Rifai, M. A. 1968. The Australasian Pezizales in the Herbarium of the
Royal Botanic Gardens Kew. Ver. Kon. Ned. Akad. Wetensch., Afd.
Natuurk. 67:1-295.
Thind, K. S. and L. R. Batra. 1957. The Pezizaceae of the Mussoorie
hills. IV. Jour. Indian Bot. Soc. 56:428-438.
and. P. Singh. 1959. Idem, Vi. «ibid, “oe; 221-232,
and K..S. Waraitch. 1970, The Pezizales’ of India.) Xx. kes.
Bull. Panjab University 27:145-155.
, E..K. Cash and P. Singh. 1959. The Pezizaceae of the Mussoorge
Hills (India). VII. Mycologia 51:458-464.
Warditch, K. S. and K. Si'Thind. 1977. Fungi of Nepal (Pezazaies).
J. Nat. Hist.-'Mus, 1:21-34.
MYCOTAXON
VORA I. No. 1, pp.2 1234132 October-December 1982
ISOLATION AND IDENTIFICATION OF EUTYPA ARMENIACAE
FROM DISEASED GRAPEVINES IN WASHINGTON STATE
Deana. GLAWE* and C. B. SKOTLAND
Irrigated Agriculture Research and Extension Center
Washington State University
Box 30, Prosser, WA 99350
and
W. J. MOLLER“
Department of Plant Pathology
University of California, Davis, CA 95616
SUMMARY
Eutypa armeniacae is reported in Washington state from
diseased grapevines that exhibit symptoms typical of Eutypa
dieback. The teleomorph was not found. The fungus was i-
dentified on the basis of host symptomatology, formation of
a Cytosporina state in culture, and pathogenicity on apri-
cot. Problems in the taxonomy and identification of E. ar-
meniacae are discussed.
Eutypa armeniacae Hansf. & Carter ex Carter (anamorph:
Cytosporina sp.) (Pyrenomycetes, Sphaeriales, Diatrypaceae)
causes cankers and diebacks on grapevine, Vitis vinifera L.
and V. labrusca L. (Moller and Kasimatis, 1978), and apri-
cot, Prunus armeniaca L. (Carter, 1957) and has been re-
ported from a number of other woody plants. The fungus
ranks as a major pathogen of both grapevines and apricots,
and occurs throughout the world. Unfortunately, identifi-
cation of this fungus is difficult. There is no monographic
etesteys address: Department of Plant Pathology, Universi-
byoor Illinois, Urbana, LL 6LS0L,
eran ete
124
treatment of Eutypa, and taxonomic literature on the genus
is scattered and often inaccessible. In addition, the type
description of E. armeniacae (Carter, 1957) does not clear-
ly differentiate it from a number of older Eutypa species,
includifig Ea acharii1 Tul., EB. lata (Pers.)) > Fru
and E. ludibunda Sacc. (Saccardo, 1882). Thus, the dis-
tinctions between E. armeniacae and some other species are
unclear. Another difficulty is that the teleomorph of E.
armeniacae is not known to form on natural substrata in re-
gions receiving less than approximately 33 cm of annual
rainfall. or in artificial culture (Carter, 4195/7)-eeconce—
quently, in dry regions E. armeniacae must be identified on
cultural and anamorphic characters without reference to
teleomorphic characters. Although the Cytosporina state
forms in culture, the lack of available information onsGy—
tosporina states of other species of Diatrypaceae prevents
confident identification of E. armeniacae solely on ithe
morphology of its anamorph. Thus, in addition to cultural
studies, previous investigations have used its pathogenic-
ity on apricot to help differentiate E. armeniacae from
other Eutypa or Cytosporina species (Carter et al., 1964;
eee
Dingley, 1960; English and Davis, 1965; Moller, 1964;
Molier ‘et als," l9638;" Moller vetwalcy 97); sUyemotometecains
1976).
The difficulties in identifying E. armeniacae hayewor—
ten impeded research on the diseases caused by it. Such
has been the case in Washington state, where in 1975 (Mink,
1975) a grapevine disease resembling Eutypa dieback was re-
ported from the semi-arid Yakima Valley, which receives ap-
proximately 18 cm of annual rainfall (Anonymous, 1978).
Attempts to find the teleomorph of E. armeniacae failed,
and although some fungi isolated from the vines resembled
E. armeniacae in cultural characteristics, they did not
sporulate and were not identified (Mink, 1975).
This paper reports the results of subsequent efforts
to identify E. armeniacae isolates from diseased grapevines
in Washington. Identification of E. armeniacae in this
Study was based on: 1) symptomatology on grapevine; 2)
production of a Cytosporina state in culture; 3) pathogen-
icity on apricot. Problems in the taxonomy and identifica-
tion of E. armeniacae are discussed, and etiology of Eutypa
dieback of grapevine in Washington is discussed in relation
to identification of the fungus.
MATERIALS AND METHODS
Vineyard observations. Twenty-four 10-yr old or older
"Concord' (Vitis labrusca) vineyards in the Yakima Valley
125
of Washington state were surveyed in late May and early
June, 1976-1978 for presence of the disease. Disease inci-
dence was estimated in five vineyards by randomly choosing
a vineyard block in each that contained 600-1800 vines, and
examining each vine for foliar and trunk symptoms. The
same vineyard blocks were examined each year.
Isolation and cultural techniques. Diseased wood was
excised from canker margins on 60 grapevines and stored at
4C until isolations were made. Pieces of wood approximate-
ly 0.5 cm square were surface-sterilized 30-90 sec in 1.622
sodium hypochlorite, and placed on several media, including
potato-dextrose agar (PDA), PDA acidified by adding one
drop of lactic acid per 9-cm-diam Petri plate, and PDA con-
taining 200 ppm streptomycin sulfate and 50 ppm tetra-
cycline. Isolates obtained in this manner were transferred
to 9-cm-diam plastic Petri plates containing 30 ml PDA and
incubated either in darkness or on a laboratory bench where
they were subject to fluorescent room lighting. Ambient
temperatures during incubation ranged from 20-28C. Two
known isolates of E. armeniacae from California, no. 590
(isolated from apricot, Prunus armeniaca) and no. 1090 (i-
solated from grapevine, Vitis vinifera), were cultured un-
der the same conditions for comparison.
Pathogenicity tests. Tests? were ‘conducted,on apricot
to assess the pathogenicity of two suspected E. armeniacae
isolates from Washington grapevines; these tests also in-
cluded known E. armeniacae isolates no. 590 and no. 1090
from California for comparison. The Washington isolates
were no. 261, isolated from a diseased grapevine near Sun-
nyside, Yakima Co., June, 1976, and no. 783, isolated from
a diseased grapevine near Grandview, Yakima Co., June,
1977. The pathogenicity tests were conducted at Davis, CA,
and at Prosser, WA. 'Tilton' apricot trees were utilized
at Davis. Because varietal trees were unavailable at
Prosser, apricot trees that had been started from seed by
Dr. T. Toyama were used there; parentage of the Prosser
trees is unknown. Inoculum consisted of discs of mycelium
cut from PDA plates; inoculum in control treatments con-
sisted of sterile PDA discs. Inoculations were made by
pruning off lateral branches 1.0-1.5 cm diam, and placing
inoculum on the wounds. The inoculation points were then
wrapped with aluminum foil to prevent rapid desiccation.
Test results were evaluated by splitting inoculated
branches longitudinally and measuring the total lengths of
xylem discoloration in both directions from the inoculation
points. The fungi were reisolated as before. Five trees
were inoculated with each of the five treatments (four
126
isolates and control) in the Prosser test, and four trees
with each treatment in the Davis test. The Prosser test
ran from May 16-October 13, 1980, and the Davis test ran
from May 28-November 21, 1980.
RESULTS
Vineyard observations. The disease was found in each
of the 24 vineyards. Disease incidence in the five select-
ed vineyards ranged from 20-84%, and did not change signif-
icantly during 1976-1978.
Foliar symptoms were most evident in late May when
new shoots were three or four internodes long. Symptoms
became more difficult to detect as the season progressed
because diseased foliage tended to be obscured by healthy
shoots. Diseased leaves were stunted, chlorotic, and cupu-
late. Internodes were shortened and the berries tended to
fall from affected shoots. Terminals of severely diseased
vines often collapsed and died during hot weather; new
shoots often developed below the diebacks the following
spring. Unlike healthy vines, diseased vines usually suck-
ered profusely.
Diseased vines also had trunk cankers associated with
large pruning wounds that measured 5-8 cm in diameter and
had been made during the mid-1960's when trellises were
changed from four-arm Kniffen trellises to two-arm trellis-
es. Smaller wounds caused by normal pruning operations
were seldom associated with cankers. Cankers occasionally
were evinced by concentric peripheral ridges resulting from
growth of adjacent healthy tissue, but usually were incon-
Spicuous unless covering bark was stripped away to reveal
the dark-brown-discolored wood typical of the cankers. The
discolored wood usually extended below ground on vines with
severe foliar symptoms. Vines died when cankers girdled
their trunks.
A Cytosporina state resembling that of E. armeniacae
was found on the canker of one diseased vine near Prosser,
WA in June, 1977. The teleomorph was never observed.
Isolations and cultural observations. A fungus which
formed a Cytosporina state in culture was isolated from 50
of 60 randomly chosen diseased vines. Colonies were ini-
tially hyaline with cottony aerial growth and diffuse mar-
gins. After 2-4 wk, regions of dark gray to black aerial
hyphae developed. Isolates differed in the extent and in-
tensity of the dark mycelial regions. Pale yellow conidial
masses began exuding from black, 1-mm-diam, subconical pyc-
nidia after 4-6 wk. Conidia were 34-74 (-78) x 1-1.5 (-2)
um, single-celled, moderately curved, filiform, and
LZ
Table 1. Pathogenicity of four isolates of
Eutypa armeniacae on apricot (Prunus
armeniaca) at Davis, CA and Prosser, WA.
Ave. Length Internal
Discoloration (Cm)*
Isolate Davis Prosser
Control Alva is0sa
261" lyfe Thea ENG
10907 30.0 ¢ 3.3 ab
783> 34.0 ¢ 4.1 b
5907 44.8 d 7S Olec
*Values within columns followed by the same letter are
not significantly different from each other (P = 0.05)
according to Duncan's new multiple range test.
YWashington isolate.
Z ‘ ‘i A
California isolate.
hyaline. Reverse sides of month-old colonies were creamy
white to pale yellow. Cultural characteristics of the
known E. armeniacae isolates from California were identical
to Washington isolates while the colonies were expanding,
but sporulating cultures (4-6 wk old) differed in several
respects. The California isolates did not become as dark
as Washington isolates, tended to produce more pycnidia
with larger spore masses and smaller conidia that measured
(15-) 25-35 (-39) x 1-2 um, and discolored the culture me-
dium a more intense yellow. Neither the Washington nor the
California isolates sporulated in darkness.
Pathogenicity tests. Results of pathogenicity tests
of two suspected isolates of E. armeniacae from Washington
grapevines and two known E. armeniacae isolates from Cali-
fornia are presented in Table 1. All four isolates were
pathogenic in the Davis test. At Prosser, isolates no. 590
and no. 783 were pathogenic, but isolates no. 261 and no.
1090 did not differ significantly from the control treat-
ment. Xylem discoloration extending from wounds inoculated
with isolate no. 590 was significantly greater than that
produced by other isolates in both tests.
128
DISCUSSION
This=report is one of several (Carter et al., 11964.
English and™Davis, 1965; Kouyeas et al.) 1978; Kouyeas eu
al., 1976) in which the teleomorph of E. armeniacae was not
found on the host from which the fungus was reported. I-
dentification of E. armeniacae in the absence of its teleo-
morph under natural conditions is complicated by its ina-
bility to form the teleomorph in artificial culture. It is
further complicated by a general lack of information on
Cytosporina states which might be confused with the Cyto-
sporina state of E. armeniacae. Previous studies on E.
armeniacae have used its pathogenicity on apricot and be-
havior in culture to help distinguish it from other fungi
(Carter et al., 1964; Dingley, 1960; English and Davis,
£9655, Molter, (1964 "Moller etal) 1968s °Mollerscteal
LT Uy emocoveteral on. Lo7 6) Consequently, the following
three criteria were ood to identify E. armeniacae in this
study:
1) Consistent isolation of the fungus from diseased
vines exhibiting symptoms identical to those caused by E.
armeniacae in other areas (e.g., Moller and Kasimatis,
1978). Disease symptoms included stunted and deformed
chlorotic foliage, dieback, and trunk cankers associated
with pruning wounds. These symptoms are, insofar as is
known, unique to Eutypa dieback.
Zy Ve Productionvor <a Cytosporina state in artificial
culture when exposed to light. Although some other species
of Diatrypaceae sporulate in darkness (Glawe, unpublished) ,
E. armeniacae does not. Conidia of Washington isolates are
larger than the size range reported in the type description
(Carter, 1957) and found in the known E. armeniacae iso-
lates examined in this study. However, R. C. Pearson (per-
sonal communication) reports that ascospore isolates from
grapevines in New York produce conidia that are similar in
size to Washington isolates. Thus, it appears that the
original description of E. armeniacae does not indicate the
full size range of conidia formed by this species.
3) Pathogenicity on apricot. Apricots were used in
this, as well as in previous studies (Carter et al., 1964;
Dingley, 1960; English and Davis, 1965; Moller, 1964; Moll-
Cra ctgale 1966; Moller et 'al.’,. 1968; Moller etvala, ta.
Uyemoto | et alate 60) to verify RA ealosees of isolates
from diseased apenas Apricots are preferred in patho-
genicity tests for purposes of identification because symp-
toms develop in months rather than years as is required on
grapevines (Moller and Kasimatis, 1978). Both the known
129
isolates (no. 590 and no. 1090) and the suspect isolates
(no. 261 and no. 783) were pathogenic in the Davis test.
In the Prosser test, isolates no. 590 and no. 783 were
pathogenic, but isolates no. 261 and no. 1090 did not dif-
fer significantly from the control treatment. From these
results it appears that the latter isolates were less viru-
lent, and might have become differentiated from the control
treatment in the Prosser test had that test continued long-
er. Differences in virulence among E. armeniacae isolates
have been reported previously (Ramos et al., 1975a). Iso-
late no. 590 caused significantly more xylem discoloration
than the other isolates in both tests. Discoloration of
xylem was more extensive for each isolate, and in the con-
trol treatments, in the Davis test than in the Prosser
test. The reason for this is unknown. The Davis test ran
approximately six months and the Prosser test approximately
five months, but it appears unlikely that the difference in
test lengths is responsible for the four- to nine-fold dif-
ferences between test results. Although no differences in
varietal susceptibility to E. armeniacae have been report-
ed, genetic differences among apricots might have affected
the test results. Another possibility is that climatic
differences between Davis and Prosser influenced the re-
sults. This possiblility is consistent with the fact that
Eutypa dieback has not been reported on apricot in Washing-
ton or other cool areas of the world where they are grown,
but is common on apricot in California and other areas with
mild climates (Moller, unpublished). Further studies are
necessary to clarify the factors affecting development of
Eutypa dieback on apricot, but the pathogenicity tests in
this study do demonstrate the pathogenic potential of Wash-
ington isolates on apricot.
Identification of E. armeniacae on the basis of host
Symptomatology, formation of a Cytosporina state in culture
and pathogenicity on apricot is consistent with previous
interpetations of this-species (Carter et al., 1964; Ding-
ley, 1960; English and Davis, 1965; Moller, 1964; Moller et
ieee Ob. Mol Verietval., 11 960s4Mol lem eteale.. b9/ ts
Uyemoto et al., 1976). However, as stated earlier herein,
the type description for E. armeniacae (Carter, 1957) is
very similar to those of a number of nomenclaturally older
Eutypa species. The description for E. armeniacae (Carter,
1957) does not indicate how it may be distinguished from
those species, nor have any subsequent studies addressed
this problem. Thus, it is possible that E. armeniacae is a
Synonym of an older name; the name is used here in order to
conform with its widespread use among plant pathologists.
130
Clearly, however, studies on E. armeniacae and similar
Eutypa species are needed to clarify their taxonomy and
nomenclature, and to simplify their identification.
The results of this study establish the presence of
E. armeniacae and Eutypa dieback of grapevine in Washing-
ton. Previous investigations of Eutypa dieback in Washing-
ton (e.g., Mink, 1975) failed to elucidate its cause, ap-
parently because fungi isolated from diseased vines were
cultured in darkness and isolates of E. armeniacae escaped
attention because the cultures failed to sporulate. The
source of inoculum in Washington remains to be determined.
Conidia of E. armeniacae are thought not to function as
inoculum because attempts to germinate them have failed
(Moller and Kasimatis, 1978). The teleomorph was not found
in the Yakima Valley, which receives approximately 18 cm of
annual rainfall (Anonymous, 1978). Areas receiving more
than 33 cm of rainfall, the amount reported as necessary
for formation of the teleomorph (Carter, 1957), are located
50 to 100 km to the west and southwest of the Yakima Valley
(Anonymous, 1978), and contain known host species. These
areas are within the 60-km range for ascospore dispersal
reported in California (Ramos et al., 1975b), and the 160-
km range reported in Australia (Carter, 1957). It appears,
therefore, that the teleomorph of E. armeniacae might occur
in these neighboring higher-rainfall areas, and that long-
range ascospore dispersal may be an important feature in
the biology of this fungus in Washington. Further research
is needed to determine the types and sources of inoculum in
Washington.
ACKNOWLEDGMENTS
Scientific Paper No. 5679, Washington State University
College of Agriculture Research Center, Project 1628. This
project supported in part by funds from the Washington Con-
cord Grape Research Council. We thank Dr. T. Toyama, Ir-
rigated Agriculture Research and Extension Center, Prosser,
for apricot trees used in this study, Dr. T. Russell, Wash-
ington State University, for assistance with statistical
analysis of data, and Dr. R. C. Pearson, Geneva Experiment
Station, NY, for making available unpublished information
regarding E. armeniacae. We thank Dr. M. E. Barr Bigelow,
University of Massachusetts, and Drs. R. Duran, J. D.
Rogers, and C. G. Shaw, Washington State University, for
reviews of the manuscript. We are especially grateful to
Drs. Duran and Rogers for valuable discussions regarding
this research, and to G. H. Glawe for helpful comments re-
garding the manuscript.
ti
LITERATURE CITED
ANONYMOUS. 1978. Climatological data, Washington. U. S.
Environ. Data Inform. Serv. 82:4-5.
CARTER, M. V. 1957. Eutypa armeniacae Hansf. & Carter sp.
nov., an airborne vascular pathogen of Prunus armen-
iaca L. in Southern Australia. Austral. J. Bot.
Sha Fs er
CARTER. DM.» V.,. G. S;:-MORVAN, and C.. CASTELAIN.). 1964. . An
extension of the known distribution of Eutypa armen-
facae. Nature 202:1134-1135.
DINGLEY, J. M. 1960. Eutypa canker of apricots. Orchard.
New Zealand 33:78-79.
ENGCLOH en. and J... Re oDAVIS. 1965. “Apricot, dieback ‘fun-
gus found on western choke-cherry. Pl. Dis. Rep.
49217 5:
KOUYEAS, H. 1978. Eutypa armeniacae on lemon in Greece.
Piyuopathol. Ze) (ols2354237.
KOUYEAS, H., A. CHITZANIDIS, A. PAPPAS, and M. V. CARTER.
1976. Eutypa armeniacae on apricot and grapevine in
Greece. Phytopathol. Z. 87:260-263.
MINK, G. E. 1975. A dying arm-like disease of grapevines.
Proc. Washington State Grape Soc. 1975:25-26.
MOLLER, W. J. 1964. Apricot disease found on garden
Srube io. eAlonralng ss Agric. 675251.
MOGEER WW. a. BH. SENGUISH, and. J. Rs DAVIS. © -1966..? “The
perithecial stage of Eutypa armeniacae in California.
Pit Diss Rep. "50353.
MOLLER, W. J., H. ENGLISH, and J. R. DAVIS. 1968. Eutypa
armeniacae on grape in California. Pl. Dis. Rep.
be <7 Len ea
MOLLER, W. J., and A. N. KASIMATIS. 1978. Dieback of
grapevines caused by Eutypa armeniacae. Pl. Dis.
Rep. 62:254-258.
12
MOLLER, Woods 50D.) B RAMOS ,- and: Ws Re eHibDRe Tie see Gaye
Apricot pathogen associated with Ceanothus limb die-
back in Calitornia. Pl. Dis. ‘Rep:, “55-2006=100c7
RAMOS 5) D. E., W. J. MOLLER, and®H. ENGLISH. ~19/5as sous
ceptibility of apricot pruning wounds to infection
by Eutypa armeniacae. Phytopathology 65:1359-1364.
RAMOS, D. E., W. J. MOLLER, and H. ENGLISH. 1975b. Pro-
duction and dispersal of ascospores of Eutypa armen-
iacae in California. Phytopathology 65:1364-1371.
SACCARDO, P. A. 1882. Sylloge fungorum omnium hucusque
Gognitorum.y) VOL... “Padua.
UYEMOTO. J. Ka, Wo J. MOLLER, and A. CC. -GOHEEN =) 1976.
Isolation of Eutypa armeniacae from grapevines and
inoculation to apricot. Pl. Dis. Rep. 60:684-666.
MYCOTAXON
Vole AVIS No. 1, pp. 1335-140 October-December 1982
STILBELLACEOUS FUNGI 1. DIDYMOSTILBE
J. Le. CRANE
Illinois Natural History Survey and the Departments of
Botany and Plant Pathology, University of Illinois, Urbana,
Illinois 61801
and
ADRIANNA D. HEWINGS
Department of Plant Pathology, University of Illinois,
Urbana, Illinois 61801
The genus Didymostilbe was established by Hennings
(1902) with D. coffeae as the type species. Twenty-six
days later Didymostilbe Bresadola and Saccardo was
described in Saccardo (1902) and typified by D.
eichleriana. Sydow (1903) noted this nomenclatural problem
and indicated that Hennings” name has priority.
We have examined the type material of six of the eight
species presently place in Didymostilbe. It appears that
this genus differs from Stilbella Lindau as lectotypified
DYA co, erythrocephala (Ditmar:Fr.) Lindau solely by the
presence of a septum in the mature conidia. This character
is highly variable and by itself, perhaps insufficient to
maintain Didymostilbe as a genus distinct from Stilbella.
In the absence of a critical examination of Stilbella, we
have retained Didymostilbe. A revision of the genus is
presented here.
Didymostilbe P. Hennings, Hedwigia 41:148. 5 August 1902
Species Typica: D. coffeae P. Henn.
134
——=
———=—
i:
Or!
SH ASe
igQes>
Ee
Se:
- Didymostilbe coffeae. A.
. Conidiogenous cells; C.
—}S
——————
=e
SS
ee
Synnema 3
———
——— —— ———=
——_ ———————
A=L—_L_$< $< = —$—S>
———
=
‘e3S2
a
ye
OC
SINCE,
Re
Gt
BSAt
<5,
ISS
= Didymostilbe Bres. & Sacc.,
Compt. Rend. Congr. Bot. Palermo 1902:59. 31
August 1902.
Species typica: D. eichleriana Bres. & Sacc.
Conidiophores macronematous, synnematous. Synnemata
erect, straight, white, cream or tan, cylindrical to
clavate, smooth with a mucoid head. Filaments of synnemata
interwoven, hyaline, septate, monopodial, dichotomous or
verticillate. Conidiogenous cells enteroblastic phialidic,
integrated or discrete, terminal or lateral, hyaline,
smooth-walled, cylindrical. Phialoconidia acrogenous ,
hyaline, 1-septate.
Key to the Species of Didymostilbe Henn. *
1. ConvaLopenousmcel ls << COMM LON sss. esicoscesceecies 2
VOMpororenousuceillsl > 20 tim LONE: .. sss ss ss sie's's oes 0105
2% COnidiavesc—4+. eume wide... e's os Disisisisieieles ee ee Lets
Conidia 422-6.6 pm wide ........... D. coprophila (2)
Creme cai diar MOM long 22. <e'o5 es scee ees De Kamatii (3)
COT Om ae MOmiMe LONE .6.0< ssles sb eects Soe sees ae eee. 4
4, COMMGMaArTUSOTU! cus cl ses sees o's s sioisha es tieeie oy MDs BOO, Leae
Gol Ms PODOVOLO Giese’ cate sss sie ies es De even) en.and
#Numbers in parentheses refer to literature cited.
Didymostilbe coffeae P. Henn. Figure 1, A-C.
Hedwigia 41:148. 5 August 1902.
Mycelium immersed in the substrate, composed of
branched, septate, subhyaline or light brown hyphae.
Conidiophores macronematous, synnematous. Synnemata erect,
straight cream or tan-brown, cylindrical, smooth, with a
mucoid head, up to 880 um high, 60-110 m wide at the base.
Filaments of synnemata interwoven, slender, hyaline,
monopodial or with dichotomous branching at the apex.
Conidiogenous cells enteroblastic phialidic, integrated or
discrete, terminal or lateral, cylindrical, hyaline,
smooth-walled, 29-35 x 3.5-4.0 wm. Collarettes poorly
developed or lacking. Phialoconidia acrogenous, hyaline,
smooth, fusoid, straight or slightly allantoid, 1-septate
at maturity, 16.5-20.6 x 3.0 4.0 im.
136
Way
, ee
AAW
\\
ay
NN
WAN \
Ny
Bf
42-4, ‘
A? 44,44 ht A N R
z 2G LV Tid YK \
Lon PAARL MWY) ts
citi" ‘f Nal We i) JAN AS
PND KU NADER \ CR
mye) ifs st PLOY \
| RY ahh v Ay)
og i
\
i
Figure 2, A-C. Didymostilbe eichleriana. A. Synnema;
B. Conidia; C. Conidiogenous cells and conidia.
Tod:
Etymology: After the host genus Coffea.
Type: Auf Zweigen von Coffea arabica L. Buitenzorg
{Bogor}, Java, A. Zimmermann, (B).
Other Material Examined: On Coffea sp., Camargo Chih,
Mexico, 28 April 1967, H. R. Conway. Intercepted at El
Paso, No. 68952 (BPI).
Didymostilbe eichleriana Bres. & Sacc. Figure 2, A-C.
Compt. Rend. Congr. Botan. Palermo 1902:59. 31
August 1902.
= Didymostilbe obovoidea Matsushima, Icones microfungorum
a Matsushima lectorum: 60. 1975.
Mycelium immersed in the substrate, composed of
branched, septate, hyaline hyphae. Conidiophores
macronematous, synnematous. Synnemata erect, straight,
white to cream colored, cylindrical to clavate, smooth with
mucoid, tan-brown heads, 500-750 x 120-130 um. Filaments
of synnemata interwoven, slender, hyaline with monopodial,
dichotomous or verticillate branching at apex.
Conidiogenous cells enteroblastic, phialidic, discrete,
terminal or lateral, cylindrical, hyaline, smooth-walled,
with a collarette, 25-50 (-55) x 1.2-2.2 um. Phialoconidia
acrogenous, collecting in tan-brown droplets, obovoid,
hyaline, smooth, 1-septate at maturity, (8.9-) 11.0-15.0 x
2.e= Se) WM.
Etymology: Honoring the collector, B. Eichler.
Type: Supra algas vivas in truncis Betulae albae in Polonia
rossica, B. Eichler, (PAD).
Didymostilbe ellisii Saxena & Mukerji Figure 3, A-E.
Trans. Brit. Mycol-Soc. 55:503. 1970.
Mycelium superficial to immersed in substrate,
composed of branched, septate, hyaline hyaphae.
Conidiophores macronematous, synnematous. Synnemata erect,
straight, white to yellow-white, smooth, cylindrical at
base gradually expanding towards the apex into a_ spherical
or subspherical head, 480-622 um high, 35-45 um wide at the
base, 65-85 um wide at the apex. Filaments of synnemata
hyaline, septate, parallel at the base, divergent at the
138
apex, simple or once or twice branched. Conidiogenous
cells enteroblastic, phialidic, integrated or discrete,
terminal or lateral, hyaline, smooth-walled, cylindrical to
somewhat flask-shaped, 9-14 x 2.2-3.3 um. Phialoconidia
formed in basipetal chains, oblong to fusoid, occasionally
curved, hyaline, thick-walled, 1-septate at maturity, not
constricted, 13-23 x 2.2-4.4 um.
Etymology: Honoring M. B. Ellis.
Type: On rabbit dung, Delhi, India, April 1969, Leg. A.
S. Saxena, DU/KS100, 1M1(143075).
Figure 3, A-C. Didymostilbe ellisii. A & B. Synnemata;
C. & D. Conidiophores, conidiogenous cells and conidia;
E. Mature conidia.
Poo
Nomina Excludendae
Didymostilbe coprophila Mirza & Qureshi, fTrans. Brit.
Mycol Soc. 54: 148. 1970. Type material not seen.
Didymostilbe kamatii Pawar & Kulkarni, Current Science
41:73-74. 1972. (AMH!) No fertile heads remain on type
specimen.
Didymostilbe capillaceae Sacc. & Bres., Annales Mycologici
1:28. 1903. (PAD!) No ferile heads remain on type
specimen.
Didymostilbe coccinea (Massee) Saccardo, Syll. Fung.
22(2): 1446. 1913.
Basionym: Hartiella coccinea Massee, Kew Bulletin
191035. .1910. (K!)
= Calostilbella calostilbe v. Hohnel.
Didymostilbe obovoidea Matsushima, Icones microfungorum at
Matsushima lectorum: 60. 1975. Type material not seen.
From the description and illustrations, it appears
identical with D. eichleriana except for the width of
the synnemata.
Acknowledgements
We thank the curators of the following herbaria for
the loan of specimens: AMH, BPI, CMI, K, MWI, PAD; and
acknowledge Clark Rogerson and Amy Rossman for their
assistance with pertinent literature.
Literature Cited
1. Hennings, P. 1902. Fungi javanici novi a cl. Prof
Dr. Zimmermann collecti. Hedwigia 41: 140-149.
2. Mirza, J. H. 1970. Didymostilbe coprophila sp. nov.
Transactions of the British Mycological Society 54:
148-150.
Somrawar,;. lL. Ss and JU. Ke Kulkarni. 1972 A new
species of Didymostilbe from India. Current Science
41: 73-74.
140
4, Saceardo, P.A. 1902. Manipolo di micromiceti nuovi.
Compt. Rend. Congr. Bot. Palermo 1902:46-60.
5. Sydow, H. & P. 1903. Nomenklatorische Bemerkungen zu
einigen kurzlich neu beschriebenen Pilzarten. Annales
Mycologici 1:176-178.
MYCOTAXON
VOLE VigoONOs. L,epp ea L4 1-154 October-December 1982
VARIABILITY OF SPORES OF DIFFERENT BASIDIOCARPS GROWING
ON THE SAME MYCELIUM IN THE APHYLLOPHORALES
Erast Parmasto & Ilmi Parmasto
iInstttute of Zoology & Botany
202200> Tartu Eetontan sk, UScr
The dimensions of basidiospores are one of the most
important characters for distinguishing closely related
species of the Aphyllophorales. However, there are many
obstacles to reliable use of such characteristics. First-
ly, for most fungal species, including the Aphyllophora-
les, we do not have available spore measurements repre-
senting the entire species but only for some, poss-
ibly unrepresentative, specimens. Thus most descrip-
tions of new species include measurements of the spores
of the type specimen only, and few descriptions include
such measurements for a local population. In monographs
and keys the absolute range of spore dimensions is Given;
in most cases neither the number of spores measured nor
the origin of the specimens studied is indicated. In too
Many cases exactly the same dimensions are given in dif-
ferent text-books where fungi of remote areas are de-
scribed, which does not inspire confidence in such data.
Most contemporary taxonomists accept the biological
concept of the species and the rightness of the popula-
tion approach to species studies. Proceeding from this
point of view, a species is distinguished not by the
spore characteristics of the holotype specimen but by
those of different specimens of different populations of
species. Like all other phenotypic characters, the spore
measurements are variable within certain limits, typical
for the species. This variability is complicated and in-
cludes several components: variability of populations
within the species, variability of specimens, variability
of one specimen during the spore discharge season (de-
pendent and/or independent of environmental factors, es-
pecially weather conditions), diurnal variability, and
variability of individual spores in a spore print ob-
tained within a short period of time from one specimen.
In the Agaricales the variability of spores has been
studied by several authors in a reasonable number of spe-
cies (see Clemengon, 1979) but in the Aphyllophorales
only in a small number of species. Some less well-known
papers on the subject include the following.
142
E. Parmasto & I. Parmasto (1976) studied a local popu-
lation Of Polyporustrnhnigophttus Pat. in Central Asia
(USSR). Differences between the mean spore lengths of some
specimens was statistically highly significant, the range
of means being 7.54-8.73 yum.
K. Bjgrnekaer (1938) demonstrated that the mean length
of the spores of the same basidiocarp of Fomes fomentartus
(Pos) Fro yvaried ‘from:21./0 amon 1] May te isso omen
September, 1931, and from 20.7 ym on 19 April to 14,0 um
in September-November, 1932.
I. Nuss (1975) found that in seven species out of the
27 species he studied the form and/or dimensions of spores
changed statistically very significantly during the sporu-
lation period. In the same paper data were given on the
mean spore measurements of different specimens of one lo-
cal community.
I,.\Parmasto & E. Parmasto (1977). studied the variabire
ity of spore dimensions during the discharge period in
three species of Phelltnus in Estonia (USSR) and found the
difference between the mean spore lengths to be statisti-
Callyvsigniticant in “erareian? (Bonds) \Panmae ei.) von
cola Niemela and in three specimens of Ph. tgntartus (Fr.)
Quél. The difference was insignificant in three specimens
Of Frowgntarcus,
However, there is a further source of possible vari-
ability not accounted for in the above investigations, and
which has been virtually unstudied in the Aphyllophorales,
i. e. the differences which may occur between spores of
different basidiocarps originating from the same mycelium.
This variability “is not only \of taxonomic interest) for it
may also reflect the genetical diversity of different
parts of an individual mycelium. So far as we are aware,
this kind of variability has been studied only by R. Age-
rer (1975) in two groups’ of basidiocarps of iP lagelicsecy]
pha mtnuttsstma (Burt) Donk and F. kavinae (Pil.) W.B.
Cooke.
MATERIALS AND METHODS
Spores of 37 pairs, nine sets of three and one set of
four basidiocarps of eight different species growing in
nature were collected. Each set was believed to originate
from the same mycelium: as a rule, members of the same set
were growing not more than 15-25 cm from each other, and
in some cases they had coalesced. Spore samples were col-
lected from each basidiocarp as spore prints, shed over a
24 h period.
For four Cortolus species, herbarium material was
used, each collection consisting of 5-6 individual basi-
diocarps which had been collected as presumably growing
from the same mycelium.
From every sample 30 spores were measured in 2 % KOH
solutiom using an eyepiece micrometer at a magnification
143
of x 700 under the microscope MBMN-¢€, and the data were
compared using Student's t-test.
RESULTS
Of the 36 pairs of basidiocarps, the difference in
spore length was statistically insignificant (at P > C.10)
in 23 cases, significant (at P < 0.05) in seven cases and
statistically-highly significant (at P < 0.001), in six
cases (Table 1). Amongst the species studied there were
some pairs of basidiocarps with a significant difference
in spore length except Fomes fomentartus, Phelltnus ptnt
(only one pair studied) and Ph. populteola (three pairs
studied). ;
Of the nine sets of three and four basidiocarps (Table
2), the difference in spore length was in only two cases
statistically insignificant between all the basidiocarps
belonging to one set. In two cases (i. e. Phelltnus aint
and Ph. ntgricans) all three basidiocarps significantly
Giffered from each other in their mean spore length; and
in the remaining sets (altogether six) two basidiocarps
had similar spores and the third had statistically differ-
ent ones. In the one set of four there were two different
pairs of basidiocarps with similar spores.
In ali four sets of basidiocarps of Cortolus spp.
(Coeviireutus, 6c. pubescens, C. vereteolor and C. \20onatus)
there were some pilei with similar spores and some with
significantly different ones (Table 3).
Table l
DIFFERENCE BETWEEN SPORES IN PAIRS OF BASIDIOCARPS*
Species, Mean length iE Q) as Pisin iG
locality, substrate, date (um) + s.e.
Fomes fomentarius (Fr.) Fr
Eston. SSR, on Betula ly Oct 24 0.1 3516 £70706 2. gx
sp., V.09 17.8402 0.20 * 2.9602 0504 5
Phellinus alni (Bond.) Parm.
Eston. SSR, on Alnus in- 6,47. 2.02'06 5. | kKK Pei2ee 0-02 1.5
cana, 15.X.69 RAN SS OR Be ' 08520 O02
Eston. SSR, on Malus, do- G, 34 0.09 3, gkKX 1,08-2.0.01 0.9
mestica, 24.VIII.70 Ota 'Os06 : POEs 0502 %
Eston. SSR, on Sorbus Oe es 0. OF, 1.5 107 80.02 0.5
aucuparia, 16.X./0 5.81 + 0.09 : 1,06, =. 0,01 -
Significance of difference: * - significant at P<
OPUS emma e are (Ole: Ate om at P< 0,000. 1) Oy — mean
ratio of spore length and width.
Basidiocarps grown together to each other.
144
Table 1 (continued)
Species, Mean length 2 20S 2 < C
locality, substrate, time (um) + s.e.
AR entree rie, Cie Earn ae We Nee ie A, seb oo a ee Oe a ae
Phellinus alni
Eston, SSR, on Alnus in- Ge 29.7 0.05 1.3 Ins: 0502 2.0%
Cana look .09 62405270706 i heres eo econ Oy 6 ‘
Eston. SSR, on Padus ra- O.06R250 2,06 1.2 Lt? 05 02 0.7
cemosa, 3.VI.69 6618.2 0208 LO ve 0.02 2
- the same, 15.X.69 oie hea 6 PLO 2.3% We O Ste 0.02 0.4
eo Leone 109" = 002
Eston. SSR, on Sorbus 6. 84et 00209 eal 12.09 270.01 1.3
aucuparia, 16.X.70 629652 :0506 ; el ate OSOt c
Eston. SSR, on Malus do- a Ol. 0, O06 1.0 te er 0? 0.0
mestica, [3.V./1 elo 2 OSOF i 1.14 2°0.02 wi
Eston. SSR, on Sorbus 6007270209 0.1 LLL ee 0202 aes
aucuparia, 7.X.69 6. 010250508 : Lal aos 502 :
Phe hartigiie (All. &)Schn.jrrat,
Sakhalin, on Abies sac- RelO ss 70.09 5 Ake L.0622.0701 0.7
chalinensis, 23.VIII.71 Oe EO SOS 1205.2 6201 ;
Lvov Reg., on Abies al- (200-2 0206 1.2 UAE Broty = lOO | 0.9
ba, 2001x569 8.10 + 0.08 : Lease 0201 .
Lvov Reg., on Abies al- OmOS eG 7, 0.6 L204 2.0502 0
bay 21,1X.69 8.09 2 *0:06 J Be04 2) O20
Sakhalin, on Abies sac- Pe 36atnOC09 On 12068280701 0
chalinensis, 23.VIII.74 7e41 22) OSE0 aM Blo pean WE GU
Caucas. Nature Reserve, S.07 2 O207 0.3 L.0G 20,01 1.2
on Abies nordmanniana, Sarl -0S10 i VOSUEsOOL ‘
235 VedD
Phe denianius s(Pr)) uel,
Eston. SSR y onpSalix 5. 9S 2 OOF Lh, 6H LE EERO) 2 0.6
caprea, 14.V.69 6.38 = 0206 L102 O01
- the same, 27.V.69 5.84 2 0.06 Lh. Deke 1 See sae WA 1.5
6. 20820006 Peet OO L
Krasnojarsk Reg., on Sa- 5,62) t) Os07 Se Lp anaes aco bi 1.0
Tex Spey 28. VELL ae 5,92 £70.06 ¢ Lela. 0201 i
Eston. (SSR, son, Salax 6,08 a ON OF 2. Ox 1, 0990501 3.0%
pentandra, 12.V1.69 6.34. 12. 0.06 d 121500502 ‘
Eston, SSR, on Salix 5.90) O06 2. 6% Loe 0702 0.7
triandra, 15,%; 69 6.27 = 0109 : dea Hota chy ORC) i
Kamchatka, on Salix sac- See Ole Ors Ll 1 09%2 0702 1.9
Chalinensis 7 LO .Vilie 7) Pe G4t* 006 4 Pe E3r 0.02 }
Eston. SSR, on Salix Seas OL OT 0.9 Lot2ea os 0.6
caprea, 7.VIII.69 5.58 + 0.09 ; Le louse Os02 :
145
Table 1 (continued)
Species, Mean length t Cate SG, t
locality, substrate, time (um) + s.e.
Phellinus nigricans (Fr.) P. Karst.
Eston. SSR, on Betula 65/072 .0,07 L. 3eee L509" 0,02 0
pendula, 7.X.69 dads O09 : 1.09 ct0).02
Krasnojarsk Reg., on Be- 6.268 07,03 2 9x L.06g220. 0 1.2
tula pendula, 28.VIII.71 673 ust On05 5 L208) 0501, 5
Kamchatka, on Betula er- 6./6 £ 0.06 24k LOJe te Oe OL 0.8
Monti ey Loov lil. 7b 65.95 20.05 x DeOoutO Ol x
Sweden, Tarna, 6.1X.74 65 3672.0. 06 798 1.08. a0 OL ak
65557 0. 06 , Lea, OSO2 i
Belovezhskaja Pushcha 6.64 4 0.08 oe 1 ee OLOT 0.4
Nature Reserve, on Betu- 6379 20,07 5 Lees O01 ;
la pendula, 24.1X.69
Kamchatka, on Betula er- 6.62) 0-06 ta U07 22 020) 1
Manis, vis VIL STL Gl 20,07 F 1209 10'702 ;
Eston. SSR, on Betula 6:66 (4.0508 0.9 Pee O02 0.9
pendula, 7.X.69 6577 0.08 oe Le LO-2, O02
Eston. SSR, on Betula 7206 = 04,10 0.3 Wel 2003 0
pendula, 7.X.69 Te Onee OOS ‘ 12 e002
Kamchatka, on Betula er- 6x67 42° 0.06 0 106 230.01 0
Man wiesdd .V LIL L 6.67 2.0.06 06, =-.0.01
Ph. pini (Fr.) A. Ames
Krasnojarsk Reg., on Di, 30) +) 0,06 0.6 IAL peer! 0
Pinus sylvestris, Doo 470.05 . Loa OFOL
28. Vice L
Ph. populicola Niemela
Belovezhskaja Pushcha, Se 2 0209 0.7 Lalo? 0205 0.5
on Populus tremula, DO 0207 : L2t 2 0502 .
24,.1X.69
Eston. SSR, on Populus Se I ea Oo. Od 0.6 15093"'0:02 0.7
tremula, 14.V1.69 5a lS ote OL O05 i‘ 15a! 0508 ‘
Belovezhskaja Pushcha 37.00 #0208 0.4 PbS as 0.502 0
Nature Reserve, on Popu- 5e05 ape On07 Lalo 20502
lus tremula, 24.1X.69
Ph. tremulae (Bond.) Bond. & Boriss.
Krasnojarsk Reg., on Po- S07 20.07 7 5kkK Leto 0,02 1.5
pulus tremula, 28.VIII.71 5275.20.06 3 Ie2l.e, 0,02 ;
Eston. SSR, on Populus De 42725 0.08 9.5% 1203. 02.02 1.3
tremula, 7.X.69 369-22 0,08 ; ML OO
Caucasian Nat. Reserve, 559072 0.05 1.0 Peis 0.02 1.8
on Populus tremula, S44 0605 < i, kaa se0. 01 :
ZasVel D
Table 2
DIFFERENCE BETWEEN SPORES IN SETS OF THREE
OR FOUR BASIDIOCARPS
Species,
locality, substrate, date
Phellinus alni
Eston. SSR, on Malus do-
mestica, 1.1X.70
Eston. SSR, on Sorbus
aucuparia, 16.X.70
Eston. SSR, on Padus ra-
cemosa, 17,1X.70
Eston. SSR, on Sorbus
aucuparia, 16.X.71
Phellinus igniarius
Eston,” Sok, On, salix
triandra, 15.X.69
Eston. SSR, on Salix
pentandra, 12.V1.69
Eston. SSR, on Salix
caprea, 14.V1.69
Eston. SSR, on Salix
caprea, 16.V1I.69
Phellinus nigricans
Eston. SSR, on Betula
pendula, 14.V1I.69
Phellinus populicola
Eston. SSR, on Populus
tremula, 2.X.69
Mean
Gam)
Sqr 4s,
aeoo
Diaiok
Ee Leise ee
lhe dhe Ibe
[ES tae Ue
Nae lee ke? Lue
Van Uh bus
He iret lb
[es Leu? Lb
ae lEue lee
It I+ I+
It I+ I+
bes
6. Ox**
13,2***
1.16
1 6h0
1,10
isu i Ube
It Ib 1+
Ee ine lke
pete disse Uae
ue hess? ol Fu
Ute Ugo lan
fet tne Tbe
+ 14 I+
Ube “lke [=
(tees
19
=
2.0%
t1 - value of Student's t for the samples of two neighbour-
ing basidiocarps; t? - value of t for extreme means.
nificance of difference marked as in Tablel.
Sig-
Table 3
DIFFERENCES BETWEEN MEAN LENGTHS OF SPORES OF DIFFERENT
PILEI OF ONE HERBARIUM COLLECTION
(value of Student's t; significance of difference
marked as in Table 1)
Coriolus hirsutus (Fr.) Quél.
Colmection: TAATIII799.. Estonian SSR, near Tartu, ion a Log
Suen lnus 9g lutinosa,. .cOll.s.M. Saar 720.X<.1979.
Méan Length = sve. of spores: pileus’ no.,,1-= 5.71 + 0.08;
ee mee ete OS tS oF Ol 6. -0..094, 8 5— 96.16 2 0.113 See
6.95 + 0.12 pm.
Pileus 1 2 3 4
no.
5 Sarees 6.0*** Sone 6, 4***
4 3.4%** Tes 0
3 3.7 *** LO
2 b gees}
Coriolus pubescens (Fr.) Quel.
Collection: TAA 52534. Primorskij Reg., Sikhote-Alin Na-
LUne ReEServV.e; On. Salix. Ssp,., coll. E...Parmasto) 11.Ix.1976.,
Meanwienden 2 1S.e6> Of Vspores: /1/—"5.09'*t 0.092 °-' 5.21 +
Ov eo tote. OS Tem a oA ee Oe LO eb = bbe 59 0127
Ose 5c) 67 0 el 0 um,
Pileus 1 2 3 Z 5
NOre
6 4,.9*** 4, 4%** 3,6*** 3.0% esta:
5 3.4%** 2.8%** 220% e6
4 Les L2G 0
S reg Oso
2 ex
Coriolus versicolor (Fr.) Quél.
COMLEGEITON2 An «Li2/ 16. BSstonian “SSR; néar Tartu, ona log
Sraberutoassp,, Coll. “A. (Kollom 27 .Vil.ioi9.
Meanwiengtie: 76.6, of spores: 1 =-..5.13..4 0.083 2
6
= 23.52
Oe Orem Bo SO tO), 09 10d = 63.9 008 2 5 eo Get 0009" um’.
Table 3 (continued)
i. 2 3 4
no.
5 geo Or 0.8 0.6
4 Zor A begs) 0.3
3 Ze 1.2
2 029
Coriolus zonatus (Fr.) Quel.
Collection: TAA 97422. Estonian SSR, Jarvselja, ona log
of Populus tremula; coll. Iy.sParmasto’ 3.414197 3%
Mean Tength* + swe. of spores: 1 i— 5743 °2°0208 92 5—95-4.
#7 0.5097 3,=°5.64 2°0. 087194 = 5.73) 200509) Sa-7 5 comes oS
pn.
Pileus
tr 2 3 4.
no.
5 Big Deak Pc hE desis) LEO
4 2.4* Tee) Om
5 1.8 1.3
2 0.4
DISCUSSION
All the pairs or sets of basidiocarps studied were
collected without establishing the presence or absence of
dark lines, isolating different mycelia in the wood; in
most cases the substratum was a growing full-size tree,
and the data were collected a long time before the papers
written by A.D.M. Rayner and N.K. Todd about the signifi-
cance of isolating dark lines in wood were published. The
only criterion for considering the basidiocarps to belong
to one specimen was their close proximity.
As demonstrated recently, in several species of the
Aphyllophorales studied from this aspect this criterion
is not always reliable (see below). Nevertheless, in most
cases the neighbouring basidiocarps of the Phelltinus
species and Fomes fomentartus studied by us have spores
with an insignificant difference between the mean lengths
and usually also between the mean values of Q@. This may
imply that the Phelltnus species and Fomes fomentartus
have long-living mycelia, and the mycelium of an individ-
ual may be largely extended in the infected tree trunk or
log. Indeed it has been pointed out (e. g. Williams, Todd
& Rayner, 1981; Boddy & Rayner, 1982) that several fungi
appear to infect standing trees as extensive single indi-
viduals, whilst ubiquitous saprophytes such as Cortolus
verstcolor and Stereum htrsutum (Fr.) S.F. Gray often form
149
numerous individuals in felled timber (see below).
There are several possible explanations for the dif-
ferences between the spores of other neighbouring basidio-
carps:
1. The difference in the mean spore size of what is
statistically significant in a technical sense does not
necessarily mean that the difference is significant bio-
logically (cf. Simpson, Roe & Lewontin, 1960: 173-175).
This especially concerns the cases when the statistical
significance is at the level of P > 0.001. As a demonstra-
tion of such a case we may indicate one pair of basidio-
carps of Phellitnus alnt growing on Padus racemosa: the
difference between the mean spore lengths was insignifi-
CopcrOnw. JUNC IOI 2.0.20) bute statistically signiti-
cant on 15 October of the same year (P = 0.03).
2. Different parts of the same mycelium are genetical-
ly different as a result of the fusion of genetically dis-
tinctemycelia (ci. Burnett & Partington, 1957; Bresinsky
et al., 1977). However, the assumption that this phenom-
enon really exists under natural conditions has been dis-
claimed in several papers published in the last few years.
(CE. Todd & Rayner, 1980: "To date we are unaware of any
convincing evidence which demonstrates effective hetero-
caryosis between wild dikaryotic isolates". See also Esser
& Blaich, 1973 and Goldstein & Gilbertson, 1981: 177).
3. The decaying tree trunk or log is inhabited by two
or several neighbouring individual mycelia of the same
species, which are genetically different and which do not
fuse due to their heterocaryon (somatic) incompatibility.
This has been demonstrated recently in several Aphyllopho-
bales (Rayner & Todd, 1977, 1979; Todd & Rayner, 1978,
1980; Coates, Rayner & Todd, 1981).
The last-mentioned reason seems to be acceptable for
our present results. The only discrepancy which does not
fit is a pair of the basidiocarps of Phellinus alnt, which
had densely coalesced but had a highly significant differ-
ence between the mean length of spores.
The difference between the mean length of spores of
different pilei of one collection in the Cortolus species
(Table 3) are easily understandable. A.D.M. Rayner and
N.K. Todd (1977) have shown that many of the closely
crowded basidiocarps of Cortolus verstcolor on one tree
stump belong to genetically different isolated mycelia.
Obviously the same is also charactristic of Cortolus htr-
sutus, C. pubescens amd C. zonatus. The mean spore length
data only are not sufficient to say exactly how many indi-
vidual specimens we had in our cases mixed in one collec-
tion, but obviously there were no less than two or three.
R. Agerer (1975) compared the mean length of spores in
two groups of the basidiocarps of Flagelloscypha mtnutts-
stma and F. kavtnae. He found that in both groups (spe-
cies) there were some basidiocarps which probably belonged
to different individuals, and that the groups of basidio-
carps were consequently mixed populations (Mischpopulati-
onen).
SO
In conclusion from our study and the results of others
we may assert that if not always, then in most cases the
different basidiocarps of the Aphyllophorales growing on
one mycelium have spores with an insignificant or small
(at P > 0.001) differences in their mean size and form.
Different specimens of one local population may have a
statistically highly significant difference in spore size,
and in some species having small basidiocarps they may
happen to be kept in a herbarium as one collection.
ACKNOWLEDGEMENTS
The authors are greatly indebted to Drs. H. Clémengon
and A.D.M. Rayner for reviewing the manuscript, for their
advice” and helpful criticism, and to Asst. Prof. G. Kivi-
magi for linguistic help.
LITERATURE CITED
Agerer, R. 1975. Flagelloscypha. Studien an cyphelloiden Basidiomyce-
ten, Sydowia 2/* 131-265, °1 Tat,
Bjérnekaer, K. 1938. Underségelser over nogle Danske Poresvampes Bio-
logy. Friesia 2 (1): 1-41, (With a summary in English: Contribu-
tions to the knowledge of the biology - particularly the spore-
discharge - of some Danish Polypores).
Boddy, L. & Rayner, A.D.M. 1982, Population structure, inter-mycelial
interactions and infection biology of Stereum gausapatum. Trans.
Briat.Mycolslo0C.)) (in press).
Bresinsky, A., Hilber, 0. & Molitoris, H.P. 1977. The genus Pleuro-—
tus as an aid for understanding the concept of species in Basi-
diomycetes, In: H. Clémengon (ed.) The species concept in Hymeno-
mycetes. Vaduz, p. 229-258,
Burnett, J.H. & Partington, M. 1957, Spatial distribution of fungal
mating-type factors. Proc. R. Physical Soc. Edinburgh 26: 61-68,
Clémengon, H. 1979, Biometrische Untersuchungen zur Variabilitat der
Basidiosporen. Sydowia, Beih. 8: 110-138.
Coates, D., Rayner, A.D.M. & Todd, N.K. 1981, Mating behaviour, my-
celial antagonism and the establishment of individuals in Stereum
hirsutum, Trans, Brit. Mycol.Soc, 76:.(1): 41-51,
Esser, K. & Blaich, R. 1973. Heterogenic incompatibility in plants
and animals, Advances in Genetics 17: 107-152,
Goldstein, D. & Gilbertson, R.L. 1981. Cultural morphology and sex-
uality of Inonotus arizonicus. Mycologia 73 (1): 167-180.
Nuss, I. 1975, Zur Okologie der Porlinge. Untersuchungen uber die
Sporulation einiger Porlinge und die an ihnen gefundenen Kafer-
arten. Bibliotheca Mycologica 45, Vaduz.
Parmasto, E. & Parmasto, I. 1976, Izmenchivost' razmerov spor u
Polyporus rhizophilus Pat. In: Materialy VI konferencij po sporo-
vym rastenijam Srednej Azii i Kazakhstana, Dushanbe, p. 219-220.
(In Russian.)
Parmasto, I. & Parmasto, E, 1977, Variation in the dimensions of
spores of Phellinus sp, sp. during the discharge period, In:
Ekologicheskie osobennosti nizshikh rastenij Sovetskoj Pribaltiki.
Vilnius, p. 183-184, (In Russian, with an English summary.)
bSd
Rayner, A.D.M. 1976. Dematiaceous Hyphomycetes and narrow dark
zones in decaying wood. Trans. Brit. Mycol. Soc. 67: 546-549.
Rayner, A.D.M. & Todd, N.K. 1977, Intraspecific antagonism in natu-
ral populations of wood-decaying Basidiomycetes. J. Gen. Micro-
pio, 103° 85-90.
Rayner, A.D.M. & Todd, N.K. 1979, Population and community structure
and dynamics cf fungi in decaying wood. Advances Bot. Res.
7: 333-420,
Simpson, G.G., Roe, A. & Lewontin, R.C. 1960. Quantitative zoology.
Revised Ed. New York, Burlingame,
Todd, N.K. & Rayner, A.D.M. 1978, Genetic structure of a natural
population of Coriolus versicolor (L. ex Fr.) Quél. Genet. Res.,
Cambio 02 $7'55-65, 1 pls
Todd, N.K. & Rayner, A.D.M. 1980, Fungal individualism, Sci. Progr.,
Oxf. 66 (263): 331-354,
Williams, E.N.D., Todd, N.K. & Rayner, A.D.M. 1981. Spatial develop-
ment of populations of Coriolus versicolor, New Phytol. 89:
307-320.
MYCOTAXON
Volo XV peNox Lipps 15 2=156 October-December 1982
A NEW SPECIES OF ENDOPHRAGMIELLA FROM
SCLEROTIA OF SCLEROTINIA MINOR
Michael T. Dunnl
Department of Botany
University of Maryland
College Park, MD, USA, 20742
ABSTRACT
A new dematiaceous hyphomycete, Endophragmiella
constricta, isolated from sclerotia of Sclerotinia minor
buried’ im an agricultural field: jin Beltsville. (NOR wr.
described and illustrated.
INTRODUCTION
An interesting fungus was observed on sclerotia of
Sclerotinia minor Jagger that was used as bait in a field
experiment Adams, and. ‘Ayers. 1982). This dematiaceous
hyphomycete developed on _ sclerotia placed’ in moist
chambers, and when isolated grew very slowly on culture
media. Although this species did not show mycoparasitic
ability AgGainstwoe mI Nor iNny Vitro eal te was quite unusual and
is herein described as a new species of the genus
Endophragmiella.
Abbreviations of herbaria follow those in the Index
Herbariorum (Holmgren et al., 1981).
TAXONOMIC PART
Endophragmiella constricta Dunn sp. nov.
Coloniae in agaro ad 21 C lente crescentes, restrictae,
effusae, fuscae. Mycelium ex hyphis ramosis, septatis,
brunneis, 2-0-2.3 wm crassis compositum. Conidiophora
macronemata, simplicia, erecta, fusca, crassitunicata,
laevia, usque ad 4 cellularia, 10.0-15.0 X Ser) Uns
Cellulae conidiogenae monoblasticae, 6.9-8.0 X 3.5 um,
cellula penultima percurrenter prolifera post conidia
successiva parta, proliferae usque ad 280 ym _ longas.
Secessio conidiarum rhexolytica, cum fimbria basali
excentrica, 1.0 X 2.0 um. Conidia fusca, crassitunicata,
laevia, oblonga vel oblongo-ellipsoidea, 2-septata, 15.0-
23.0. X 6.9-10.4 um, cellula .basali pallida, cellula
centrali constricta.
1 Address reprint requests to: Soilborne Diseases Lab., USDA, BARC-
WEST, Beltsville, MD, USA, 20705
Isolatus e sclerotio Sclerotiniae minoris, e terra,
Beltsville, Maryland, M.T. Dunn (MTD 3129). Holotypus BPI,
isotypus DAOM 180189, cultura exsiccata.
COLOmies) On arti micrale medias restricted, ceffuse,
dark brown to black with little aerial mycelium, hyphae
loosely branched, septate, brown, Bollea um wide.
Conidiophores macronematous, Simple, erect, rarely
branched, brown, thick-walled, smooth, composed of up to 4
Celis: 1070-1530 X 33.5 um. Conidiogenous cells
momoblastic, 6.9=6.0 X%°3.5 jm, terminal ‘on conidiophores,
conidiogenous cell forming on each successive, percurrent
proliferation of the penultimate cell through the
conidiogenous cell and beyond the open end that formed as a
result of rhexolytic conidium secession (Figs. ARES VIG ie
As many as 29 such proliferations forming on a single
conidiophore, reaching a length of up to 280 um. Conidia
retagming a portion of the conidiogenous cell as an
eccemamnc. edasal, trill; Ws0 X 2:0 - pm. Conidia often
remaining partly attached laterally to the conidiogenous
cells giving the appearance of sympodial proliferation
CEdses fs 40.Conidta brown, thick walled.)ssmooth:. oblong
to oblong-ellipsoid, 2-septate, the lower cell paler, the
muadwe: cel constricted, 15.0-23.0 X% 6.9-10.4 um.
GoLlonvar Growth on “artificial” wmedia Ws very slow,
attaining a diameter of only 8 mm after 24 days at 21 C on
(iVPemewnich scontarned: Oro% malt extract.y “0.1% ssoytone,
O205% yeast. extract, and 2% agar. Growth was faster on
GYSI medium, with a colony diameter of 23 mm after 24
days. This medium contained 1% glucose, 0.05% yeast
Gai rnacte Oeleensoytone, 0.001%, FeClis and (2%) agar: The
addition of iron to culture media has been shown to
increase growth and sporulation of Sporidesmium scleroti-
vorum Uecker et al. [=Teratosperma sclerotivorum (Uecker,
Ayers and Adams) Hughes] another dematiaceous hyphomycete
associated with sclerotia of S. minor (W.A. Ayers, personal
communication).
Habitat: tsolated from living sclerotia of Sclerotinia
Mmnomewnicn enad «been “buried in “an “agricultural field,
Beltsville, MD.
Specimens examined: MID 3129, June 1, 1981, dried culture
im, BPI s(holotype) and DAOM (180189, isotype); living
culture of the type deposited at ATCC 46749.
Endophragmiella constricta was isolated from a
SclerOllMuimrOter cc Lerotinia minor Jagger ‘during a ' field
experiment on the ability of Teratosperma sclerotivorum to
reduce lettuce drop caused. by S. minor. Sclerotia were
sieved from the soil and placed in petri plates with moist
filter paper; E. constricta grew and sporulated on a
sclerotium, and subsequently spread onto the filter paper.
Sutton (197,3;) ta est described the genus
Endophragmiella to include E. pallescens and E.
Canadensis, both of which were hyperparasitic on carbon-
aceous stromata of ascomycetes. Hughes (1979) redefined
154
iG
and
Fidis
dium
lea
coni
ium
d
jophores showing
10 um
1ng
Develop
Bar
(
icta
2
constr
Fig
ae
d
dophragmiella
coni
En
and
-5
iophores
]
i
10S.
F
Con
secession
conti
after
cell
1g
iogenous
d
ith phlox
conti
into
d w
ion
Sta
proliferati
arrow
F
Young con
3
F
1ne.
1ne
1%
(
=
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Sit
ome}
So
tS
oOo
~~ OU
c%
ome
—
nw
nO
voc
Ow
oc
no
S
= (ek
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ee
i
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(Syae:
= ae
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nea
ro)
Soa
ro)
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ie 2S
sek.
as!
(=
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Me
y 2
D
ag
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n ro)
ee
<- wv
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oe
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AO
eo e
er
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15S
the genus for species with rhexolytic conidium secession
followed by a regular percurrent proliferation each arising
from the cell below succesive terminal conidiogenous cells.
Endophragmiella constricta is in the group of species with
2-septate conidia which includes E. ontariensis Hughes, E.
Cisne nugnes, b< trnipartita,mugnes, /E. biseptata (Peck)
Hughes and E. hughesii Hawksworth. None of these species
has been grown ge cCUsbik ae (OSeed ie Hughes, personal
communication). A recently described species, E. fallacia
Kirk (1981), also has 2-septate conidia but they are larger
than those of [3 CONnSUniCia and CyIANar Teal.
Enappnregniellas constricta 1s. distinct. from ‘the , other
species in this group in the smaller size and constricted
Shape of the conidia.
Endophragmiella constricta shares with other species
of the genus the characteristic method of percurrent pro-
liferation. Often the conidium remains associated with, or
partially attached to, the wall of the conidiogenous cell
and after several conidia have been produced, it appears as
if conidia had developed sympodially (Figs. 1, 4). Kirk
(1982) reported the characteristic percurrent type of
DRoulTerationiin FE. corticota Kirk but when the conidium
failed to secede proliferation proceeded laterally through
the wall of the conidiogenous cell. Proliferation can also
appear sympodial in E. hymenochaeticola Hughes because the
apex of the conidiogenous’ cell iS” LCONSerrceec ) and
proliferation may take place through the lateral wall
instead of percurrently (Hughes, 1978).
Endophragmiella constricta has not shown any
MYcOPaArositnceaDIity In Vitro against sclerotia of S$.
minor. Another dematiaceous hyphomycete recently described
fROnesOLerovla ws sArthrocristula hyphenata, Sigler,..Dunn, .and
Carmichael, apparently has no mycoparasitic ability (Sigler
CUe ele lUO7).. Other dematiaceous hyphomycetes that have
beehayisoratea trom. sclerotia of S.. minor ‘with proven
mycoparasitic ability include Teratosperma sclerotivorum
(Uecker sete al... 19/8; Hughes, 1979) “and T. oligocladum
Uecker, Ayers and Adams (1980). These two species grow
poorly on standard culture media but develop well on living
sclerotia of Sclerotinia species. Another morphologically
Similar hyphomycete, Laterispora brevirama Uecker, Ayers
and Adams (1982), was isolated from sclerotia of S. minor
but colonized sclerotia only in association with the above
mentioned Teratosperma spp. Evidence indicates that these
three species are biotrophic parasites with facultative
saprophytic ability (Ayers and Adams, 1979, 1981; Uecker et
al.-, 1982). Endophragmiella constricta and A. hyphenata
seem to be more opportunistic, possibly functioning as
secondary colonizers of the sclerotia.
ACKNOWLEDGEMENTS
The author would like to thank S.J. Hughes and J.L. Crane for
critically reviewing the manuscript, and D.P. Rogers for correcting
the latin diagnosis. The author would also like to thank S.M. Sherren
and P.B. Adams for first noticing this fungus and bringing it to the
author's attention. This work was carried out while the author was in
Ist:
a cooperative research associate program between the University of
Maryland and the Soilborne Diseases Laboratory, USDA, Beltsville, MD;
grateful appreciation is expressed to the latter institution for
providing facilities and funding.
LITERATURE CITED
Adams, P. B. and W. A. Ayers. 1982. Biological control of Sclerotinia
lettuce drop in the field by Sporidesmium sclerotivorum.
Phytopathology 72:485-488.
Ayers, W. A. and P. B. Adams. 1979. Mycoparasitism of sclerotia of
Sclerotinia and Sclerotium species by Sporidesmium sclerotivorum.
Canadead.. Microbiole 2521) -23.
Ayers, W. A. and P. B. Adams. 1981. Mycoparasitism of sclerotial
fungi by Teratosperma oligocladum. Canad. J. Microbiol.
27:886-892.
Holmgren, P. K., W. Keuken and E. K. Schofield. 1981. Index
Herbariorum. I. The herbaria of the world. 7th ed. Regnum Veg.
106:1-452.
Hughes, S. J: 1978. New Zealand fungi 25. Miscellaneous species. New
Zealand J. Bot. 16:311-370.
Hughes, S. J. 1979. Relocation of species of Endophragmia auct. with
notes on relevant generic’ names. New Zealand J. Bot.
17:139-188.
Kirk, P. M. 1981. New or interesting microfungi II. Dematiaceous
hyphomycetes from Esher Common, Surrey. Trans. Brit. Mycol. Soc.
77:279-297.
Kirk, P. M. 1982. New or interesting microfungi IV. Dematiaceous
hyphomycetes from Devon. Trans. Brit. Mycol. Soc. 78:55-74.
Sigler, L., M. T. Dunn and J. W. Carmichael. 1982. Arthrocristula and
Arthropsis, two new hyphomycetes with dematiaceous arthroconidia.
Mycotaxon 15:409-419
SULTONG eb.cet. 1973. Hyphomycetes from Manitoba and Saskatchewan,
Canada. Mycol. Pap. 132:1-143.
Uecker, F. A., W. A. Ayers and P. B. Adams. 1978. A new hyphomycete
on sclerotia of Sclerotinia sclerotiorum. Mycotaxon 7:275-282.
Uecker, JheuAes ows tase Ayers .and) P...b. eAdams. 1980. Teratosperma
oligocladum, a new hyphomycetous mycoparasite on sclerotia of
Sclerotinia sclerotiorum, S. trifoliorum, and S. minor. Mycotaxon
10:421-427.
Uecker. Rey Asa Wel tle enyers ‘and P. B. Adams: 1982. Laterispora
brevirama, a new hyphomycete on sclerotia of Sclerotinia minor.
Mycotaxon 14:491-496.
MYCOTAXON
Voie xvi. NO. 1,7 pp. 157-16) October-December 1982
THREE NEW SPECIES OF HYPOGYMNIA FROM WESTERN
NORTH AMERICA (LICHENES: HYPOGYMNIACEAE)
Lawrence H. Pike
Department of Botany, Oregon State University
Corvallis, OR 97331 U.S.A.
and
Mason E. Hale, Jr.
Department of Botany, Smithsonian Institution
Washington, D.C. 20560 U.S.A.
Abstract:--Two new species in the Hypogymnia enteromor-
pha group, H. heterophylla Pike and H. occidentalis Pike, and a
California endemic, H. mollis Pike and Hale, are described.
Hypogymnia heterophylla Pike, sp. nov. Fig. 10
Thallus laxe adnatus, lobis linearibus, libris, margine
nigromarginatis, medulla nigra, sorediis isidiisque destitutis; apo-
thecia numerosa; sporae ca 4 X 7 pm.
Thallus loosely attached at the base, almost subfruticose,
rather soft, whitish gray, 8-15 cm broad; lobes variable, 1-4 mm
wide but usually long and little branched, free and trailing, the
tips often perforate, the margins conspicuously black rimmed;
medullary cavity dark above and below; lower surface shiny, jet
black, rugose. Pycnidia very common; microconidia rod shaped
to weakly bifusiform, 5-6 pm long. Apothecia common, short
stipitate, 3-15 mm in diameter, the disc dark brown, plane,
splitting radially with age; hymenium 40-45 pum high; spores
8/ascus, colorless, ellipsoidal, ca 4 X 7 pm.
Chemistry: Atranorin, protocetraric acid, physodalic acid,
and physodic acid.
Type: On stunted conifers along Pygmy Forest Reserve
nature trail, Van Damme State Park, elev. 50m, Mendocino
Co., California, M. E. Hale no. 49365, 9 June 1977 (US, holotype;
duplicates to be distributed in Lichenes Americani Exsiccati).
Specimens examined (all in US). California: Del Norte Co.,
Doty 3412; Humboldt Co., Becking 61041020, Hale 56798; Marin
Co., Hermann 17540; Mendocino Co., Tucker 6089. Oregon:
158
Coos Co., Hale 48934; Curry Co., Hale 49074; Douglas Co.,
Leuthner 24432; Lane Co., Hale 48821; Tillamook Co., Pike 3144.
Hypogymnia occidentalis Pike, sp. nov. Fig. la
Thallus adnatus, lobis irregulariter inflatis, brevibus, mar-
gine concolori, medulla nigra; apothecia numerosa; sporae 5 X
7-9 pum.
Thallus rather closely adnate throughout, whitish mineral
gray, 5-10 cm broad; lobes rather short and irregularly inflated,
2-4 mm wide, lacking a black rim, the medullary cavity black-
ening above and below; lower surface black, deeply rugose,
sparsely perforate. Pycnidia abundant; microconidia rod shaped
to weakly bifusiform, 5-6 um long. Apothecia abundant, sub-
stipitate, 3-10 mm in diameter, the rim upturned, the disc dark
brown; hymenium 40-45 pum high; spores 8/ascus, colorless,
ellipsoidal, 5 X 7-9 um.
Chemistry: Atranorin and physodic acid.
Type: Big Canon, Wallowa Co., Oregon, EP. Sheldon 8763,
23 Aug. 1897 (US, holotype).
Selected specimens examined (all in US). Alaska: Flat
Island, Thomas 145. Oregon: Crook Co., Hale 49690; Curry Co.,
Hale 48921; Grant Co., Hale 49688; Union Co., Sheldon 9024.
Washington: Spokane Co., Bonser s.n. Idaho: Bonner Co., Hale
49010; Boundary Co., Hale 48236. Montana: Flathead Co., Hale
48367; Lake Co., Barkley 2004; Lincoln Co., Hale 49562;
Missoula Co., Hale 48834; Ravalli Co., Hale 48401; Sanders Co.,
Hale 48275. California: Humboldt Co., Hale 49730; Los Angeles
Co., Wheeler 1554; Mendocino Co., Hale 49340; Plumas Co.,
Hale 51971; Siskiyou Co., Hale 51627; Sonoma Co., Hale 48295;
Tehama Co., Hale 52286.
The name "enteromorpha" has been applied indiscriminate-
ly to virtually every nonsorediate Hypogymnia species in North
America at some time. The eastern population was not split
off until 1973 as H. krogii Ohlsson. Up to now the far more
varied western populations have not been critically studied.
Much of the material labeled "H. enteromorpha" in the
western states has in fact turned outto be H. imshaugii Krog, a
widespread species, notoriously variable but easily recognized by
the white medullary cavity (Krog, 1968). Almost all other
species have a dark medulla. Further breakdown and under-
standing of this latter group has depended on accurate typifica-
tion of the two oldest names, H. duplicata (Ach.) Rass. and H.
enteromorpha (Ach.) Nyl., something we have now been able to
do by examining appropriate lectotype materials in the British
Museum (Smith herbarium) and isotypes at Helsinki (Acharian
159
herbarium). Hypogymnia duplicata, a conspicuous but rather
rarely collected coastal lichen from Alaska south to Oregon, is
the species so clearly illustrated by Krog (1968, p. 95). The
usual chemistry is atranorin, protocetraric acid, physodalic acid,
and accessory diffractaic acid. We have not demonstrated
physodic acid in any specimens, concurring with Ohlsson (1973).
We have based the typification of H. enteromorpha on the
rich material in the Smith herbarium (BM) and comparison with
the scraps in H. These specimens are strongly inflated, with-
out a conspicuous black rim, more or less loosely adnate but
sometimes somewhat trailing, and produce atranorin, protoce-
traric acid, physodalic acid, physodic acid, and accessory dif-
fractaic acid (misidentified as barbatic acid by Krog using the
microcrystal tests). In addition there is a second large popula-
tion, also represented on the lectotype sheets, an acid-free
chemotype containing only atranorin, which is morphologically
indistinguishable from P+ plants. This species complex is very
common from Alaska south to California, generally toward the
coast in the Cascade ranges but also extending inland as far
east as Idaho, where a few other typically Cascade lichens, such
as Platismatia stenophylla and Pseudocyphellaria anomala, have
been found.
The first of our new species, H. heterophylla, differs from
H. enteromorpha in these respects: Lobes long and trailing, not
grossly inflated, attached mostly at the center below, margins
with a broad black zone, spores ellipsoidal (in H. enteromorpha
nearly spherical, 5 X 6 pm), and diffractaic acid always lack-
ing. Furthermore it has a much more restricted habitat and
range, pine barrens, Pinus contorta in sand dune areas, Quercus
wislizenii chaparral, etc., all within a few kilometers of the
Pacific coast from central California to Oregon.
The second new species, H. occidentalis, is widespread in
the Cascades (one collection from Alaska) and especially the
northern Rocky Mountains in the United States, overlapping to a
large extent—excepting Montana—the range of H. enteromorpha.
It often grows on lower boles of Douglas fir and other conifers
in dense forests and has a broad altitudinal range, from sea
level to 1500 m. It has fairly distinctive morphological and
chemical characters. The lobes are moderately inflated with-
out a strong (or with no) black margin, as in H. enteromorpha,
but more closely adnate with no tendency for the lobes to
trail. The chemistry, atranorin and physodic acid, is another
good diagnostic character, useful in separating the species not
only from typical P+ H. enteromorpha but also from its P-
(atranorin only) chemotype.
160
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161
Hypogymnia mollis Pike and Hale, sp. nov. Bigatti
Thallus arcte adnatus, mollis et fragilis, lobis brevibus,
congestis, superne diffuse sorediatis; apothecia ignota.
Thallus closely adnate on twigs, soft and fragile, white to
grayish white, 3-6 cm broad; lobes short, about | cm long, and
crowded, 1-3 mm wide, little branched, the upper surface at
first continuous, shiny, but soon rugose, cracking and flaking and
becoming diffusely sorediate over the whole surface; medullary
cavity brown to blackening above and below; lower surface dull
black, rugose, the tips perforate, part of the lower cortex often
eroding away to expose the medulla. Apothecia and pycnidia
not seen.
Chemistry: Atranorin and physodic acid.
Type: On shrubs in sandy area at Los Osos Oak Reserve,
near Los Osos, San Luis Obispo Co., California, elev. I5 m, M.E.
Hale no. 57768, 31 July 1980 (US, holotype; to be distributed in
Lichenes Americani Exsiccati).
Specimens examined. California: San Diego Co., Cota
1447b (US), Du Rietz 206 (USP, US).
This rare but unusual Hypogymnia is easily recognized by
the laminal diffuse soralia and the P- chemistry. It has no ob-
vious relationship with any other species in North America.
Hypogymnia laminosorediata Hawks. from Morocco has the same
chemistry and extensive laminal soralia but the thallus is much
larger and the medulla white on the roof of the cavity. Sore-
diate H. tubulosa (Schaer.) Hav., better known to lichenologists,
also produces atranorin and physodic acid but has soralia de-
limited to the lobe tips. The three collections we have seen
come from open coastal chaparral or scrub.
References
Krog, H. 1968. The macrolichens of Alaska. Norsk Polarinst.
Skrifter 144:1-180.
Ohlsson, K. 1973. New and interesting macrolichens of British
Columbia. Bryologist 76:366-387.
MYCOTAXON
Vol. UAVIS. Noe Ly ipp yh l62—164 October-December 1982
A NEW SPECIES OF PARMELIA (LICHENES)
WITH PROTOCETRARIC ACID
Mason E. Hale, Jr
Department of Botany, Smithsonian Institution
Washington, D.C. 20560, U.S.A.
Abstract.--A new species, Parmelia protosulcata Hale,
is described from the Southern Hemisphere. It contains the
depsidone protocetraric acid.
The genus Parmelia is typified by P. saxatilis (L.) Ach., a
well-known boreal lichen characterized by effigurate pseudo-
cyphellae (Hale, 1981). There are about 30 closely related
species in this group, centered mostly in eastern Asia. While
the majority contain salazinic acid, we have long Known that P.
discordans Nyl. is anomalous in containing protocetraric acid
(along with lobaric acid), giving a negative KOH test in the
medulla (Culberson, 1970). A lesser known species P. pseudo-
sulcata Gyelnik, also produces this acid (Ohlsson, 1973). We can
ngw add the species described below to this interesting com-
plex, which now includes P. crambidiocarpa Zahlbr. from New
Zealand, P.kerguelensis Crombie from the subantarctic islands,
and P. discordans and P. pseudosulcata.
Parmelia protosulcata Hale, sp. nov. Figis
Thallus adnatus, corticola, lobis sublinearibus, 2-3 mm
latis, superne effigurato-pseudocyphellatis, planis, margine
sorediatis, subtus nigris, rhizinis simplicibus vel sparse furcatis.
Apothecia rara, sporis simplicibus, incoloribus, 7-9 X II-I3 jum.
Thallus adnate to loosely adnate on bark, pale greenish to
brownish mineral gray, 2-6 cm broad; lobes sublinear, short,
contiguous, 2-3 mm wide, the surface plane, rarely weakly
foveolate, becoming faintly pruinose at the tips, deeply reticu-
lately fissured in older parts; pseudocyphellae effigurate, small,
rather sparsely developed, mostly toward lobe tips or on the
margins, usually elongate; soralia developing on lobe tips or
margins, usually orbicular but fusing and linear or densely ag-
163
Fig. 1. Isotype of Parmelia protosulcata Hale in US.
gregated with age, the soredia coarse; cortex paraplectenchyma-—
tous, 14-I6 jum thick, algal layer ca I5 pm thick, medulla white,
80-110 pm thick, lower cortex carbonized, 15-18 pm thick;
lower surface black, shiny, moderately rhizinate, the rhizines
simple to sparsely furcate or weakly squarrosely branched. Apo-
thecia rare, substipitate, 4-5 mm in diameter, the amphithecium
rugose, pseudocyphellate; hymenium 60-65 pum high; spores 8,
simple, colorless, 7-9 X 11-13 pm (data from Imshaug 45613).
Chemistry: Atranorin and protocetraric acid.
Type: On Nothofagus antarctica in an open N. antarctica
forest, Cabezera Lago at eastern end of Lago Fagnano, Tierra
del Fuego, Argentina, R. Santesson 7955, 30 Mar 1940 (S, holo-
type; US, isotype).
This is a strictly austral species, very common with P.
sulcata Tayl. in the Straits of Magellan region and southern.
Chile, especially in moist Nothofagus forests. It is also present
on at least one of the subantarctic islands. The orbicular,
164
mostly marginal soralia are quite distinct from the linear, lam-
inal ones of P. sulcata, its closest relative, which is also distin-
guished by densely squarrosely branched rhizines.
Two specimens from Chile (Prov. Cautin, Rundel 7457 and
Prov. Malleco, Mahu 2474, both in US) contain fumarproto-
cetraric acid only and seem to represent a chemotype, not
entirely unexpected although this acid had not previously been
reported in the genus.
Specimens examined. New Zealand: Campbell Island,
Harris 5455 (MSC, US). Falkland Islands, Howkins 2964 (FH).
Chile: Prov. Magallanes, Imshaug and Ohlsson 43842, 43896,
44401, 44805, 44568, 45155, 45570B (all MSC), 45613 (MSC, US),
Santesson 1843 (S, US); Prov. Osorno, Imshaug 42961 (MSC);
Brunswick Peninsula, Imshaug and Harris 39095, 39260, 39328,
39329, 39404 (MSC).
References
Culberson, W. L. 1970. Parmelia discordans, lichen peu connu
d'Europe. Rev. Bryol. Lichénol. 37:183-186.
Hale, M. E. 1981. Pseudocyphellae and pored epicortex: Their
delimitation and evolutionary significance. Lichenologist
Vo)
Ohlsson, K. E. 1973. New and interesting macrolichens of
British Columbia. Bryologist 76:366-387.
MYCOTAXON
VOIR, AVI, No. Ll, spp. 165-466 October-December 1982
A new host for Sphaeronaemella helvellae
-- Pseudorhtztna sphaerospora
Donald H. Pfister
Farlow Reference Library
and Herbarium of Cryptogamic Botany,
Harvard University, 20 Divinity Ave., Cambridge, MA 02138
While examining material of Helvellaceae in North
America, a specimen of Pseudorhiztna sphaerospora (Peck)
Pouzar (= Helvella sphaerospora Peck = Gyromttra sphaero-
spora (Peck) Sacc.) was studied and found to be infected
with Sphaeronaemella helvellae (Karsten) Karsten (Pyreno-
mycetes, Melanosporaceae). Previous reports of this
fungus have been made by Seeler (1943), Wells and Kempton
(1968), Malloch (1974), and recently by Cannon and Hawks-
worth (1982). All of these reports have been based on
parasitized material of Gyromttra tnfula (Schaeffer) Quél-
et and G. anbtgua (Karsten) Harmaja. This is the first
notice of S. helvellae on P. sphaerospora. Though S. hel-
vellae does not seem to be common, it seems to be wide-
spread in North America. Wells and Kempton (1968) stated
that in the Anchorage area Gyromttra infula is frequently
infected with S. helvellae, indicating that local epidem-
ics might be quite common.
The material of S. helvellae on P. sphaerospora
differs only slightly from that on species of Gyromttra.
Seeler (1943), Malloch (1974), and Cannon and Hawksworth
(1982) described the perithecia as being superficial to
semi-immersed. The perithecia on P. sphaerospora are often
deeply embedded with only the necks projecting. In
general, the perithecia are somewhat smaller than those
described from Gyromttra spp. Perithecia from P. sphaero-
spora range from 90 to 110 um.
Virtually nothing is known of the biology of this
parasite. The presence of S. helvellae on P. sphaerospora
might indicate a close alliance of P. sphaerospora with
the species of Gyromttra but so little information is
available that no conclusion can be drawn. Certainly all
the Gyromitras and allies should be examined for this para-
site to determine if the host range of S. helvellae is
166
broader than presently understood.
Specimens examined: On Pseudorhtztna sphaerospora.
New York, Undercliff, trail to Lock Bonnie, June 27, 1899,
R. A. and A. M. Harper (FH). On Gyromitra infula. Shel-
burne, N.H. W. G. Farlow (FH, cited by Seeler). On Gyro-
mitra tnfula. In moss and on swampy ground at edge of
boreal forest beaver pond, Tarzwell, Ontario, Canada,
George P. White (965) (FH).
This work was supported by National Science Foundation
Grant DEB 8023018.
Literature Cited
Cannon, P. F. & D. L. Hawksworth, 1982. A re-evaluation of
Melanospora Corda and similar Pyrenomycetes, with a
revision of the British species. Bot. J. Linn. Soc.
84: 115-160.
Malloch, D. 1974. Sphaeronaemella helvellae. Fungi Can-
adenses, no. 53.
Seeler, E. V., Jr. 1943. Several fungicolous fungi.
Farlowia 1: 119-133.
Wells, V. and P. E. Kempton. 1968. Studies on the fleshy
fungi of Alaska. II. Mycologia 60: 888-901.
MYCOTAXON
ences ee ee ee ee Se ee ee ee ft
VOI AVIS aNOness pps 167 Sh74 October-December 1982
SMITTIUM LONGISPORUM, A NEW HARPELLALES
(TRICHOMYCETES) FROM CHIRONOMID GUTS
MARVIN C. WILLIAMS
Department of Biology
Kearney State College, Kearney, NE 68847
ROBERT W. LICHTWARDT AND STEPHEN W. PETERSON
Department of Botany
University of Kansas, Lawrence, KS 66045
The fungal genus Smittium (Harpellales, Legeriomyceta-
ceae) has 18 recognized species, including the zygospore-
producing one described in this paper. Smittium species
inhabit the hindguts of aquatic Diptera larvae such as
midges (Chironomidae, Ceratopogonidae), mosquitoes
(Culicidae), and blackflies (Simuliidae). Of the 18
species, 14 were described from midges (including the
present species), three from mosquitoes, and one from
blackflies. Some species are able to infest more than one
host family. For example, we have found S. culicis and
S. culisetae, normally in mosquito larvae, in the guts of
blackfly larvae. Williams and Lichtwardt (1972) collected
trichospores from axenic cultures of four Smittium spp.
isolated from various geographic locations and from differ-
ent dipteran hosts (chironomids, mosquito, and blackfly
larvae), fed them to mosquito larvae (Aedes aegypti), and
demonstrated that there was some degree of host preference
by the fungi at the insect family level. Zygospores have
been described from only six species, all of them from
Chironomidae. The currently known distribution for the
new species, S. longisporum, includes several midwestern
states of the U.S.A. and one site in Sweden above the
Arctic Circle.
168
"y :
169
Smittium longisporum Williams, Lichtwardt & Peterson,
sp. nov.
Trichosporae elongate fusiformi-ellipsoidales, (40-)46
(-55) x (6-)8(-10) um, collare (10-)13(-17) x ca. 4 um.
Cellulae genitales longae plerumque 1-2 trichosporae in
ramo fertili omni producentes. Zygosporae oblique affixae,
fusiformi-biconicae, (102-)110(-113) x (13-)15(-17) um,
collare 28-32 x ca. 5 um lato, ad regionem (11-)14(-18) um
a termino uno zygosporae affixo, appendiculo unico
praedito. Chironomidarum larvalium incola.
Trichospores long fusiform-ellipsoidal (40-)46(-55) x
(6-)8(-10) um with a long, well-defined appendage which
often has a zigzag appearance upon release of the tricho-
spore; collar (10-)13(-17) x ca. 4 um, slightly bulged
centrally. Usually 1-2 trichospores per fertile branch
produced on long generative cells. Zygospores fusiform-
biconical (102-)110(-113) x (13-)15(-17) um, collar 28-32
x ca. 5 um attached (11-)14(-18) um from one end of the
zygospore which bears a single appendage several times
longer than the zygospore; attached obliquely to the zygo-
sporophore which arises laterally from one of the con-
jugants. Basal cell with two downward-curved projecting
branches. Growing in the hindgut of several species of
larval Chironomidae.
Holotype: Slide DGK-1-1 deposited with R. W. Lichtwardt
at the University of Kansas, prepared from the hindgut of
a Cricotopus sp. larva collected 2 January 1981 in Rock
Creek, Section 8, T14S, R1I8E, Douglas County, Kansas, USA,
where County Road #462 intersects Rock Creek.
Figs. 1-6. Smittium longisporum. 1. Developing tricho-
spores, 575; note early collar development (subterminal
spore) followed by spore enlargement (terminal spore). 2.
Trichospores nearing maturity, x*575. 3 and 4. Mature
released trichospores with the characteristic long collar
and zigzag appendage, x650. 5. Zygospore, x535; most of
the single appendage can be seen furled within the long
collar. 6. Basal cell of a young thallus, 780; note the
branch growing from the clasplike holdfast portion of the
cell.
170
Collections: In addition to the type locality, thalli and
trichospores were found in larval Chironomidae from: a
small stream draining into the north shore of Lake
Tornetrask across from the Abisko Naturvetenskapliga
Station, Sweden, 7 July 1971; Indian Creek 3.6 km S of
highways MO 17 and US 63 junction, Texas County, Missouri,
USA, 5 February 1982; Platte River just S of highways I 80
and NE 10 (Minden interchange), Buffalo County, Nebraska,
USA, 15 April 1982; Neosho River 1.6 km E of highways
US 50 and I 35 interchange near Emporia, Lyon County,
Kansas, USA, 1 May 1982. All host collections were made
in flowing waters.
The long, relatively wide trichospores readily differ-
entiate this species of Smittium from all others presently
described (Figs. 1-4). The only species which somewhat
resembles this one is S. macrosporum Kobayasi (1969),
whose trichospores are similar in shape but are shorter
and narrower with a shorter collar, and whose walls are
described as being coarse or finely verrucose; no zygo-
spores have been reported in that species. The holdfast
of S. macrosporum was described as a short-cylindrical peg
arising from a branching basal cell, whereas in S.
longisporum the basal cell has a clasping appearance due
to the growth of two downwardly curved projections (Fig.
6), and the secreted holdfast structure present in many
trichomycetes is not evident. The basal cell is about 7
um in diam, with up to five lateral branches growing from
the downward projections in more mature thalli. The
branches of the main part of the thallus measure about
7 um in diam near the base, about 9.5 um at the widest
part, with the generative cells on terminal branches
measuring about 4 ym in diam.
Our trichospore measurements were made from phase-
contrast photographs of living material mounted in water.
We noticed that some shrinkage of trichospores (ca. 1 um
in width) occurred after the water mounts were fixed in
lactophenol-cotton blue.
Conjugation in sexual reproduction is accomplished when
two hyphae send out lateral protruberances which meet and
fuse. One of the conjugants then elongates and swells to
produce the zygosporophore, and a terminal zygospore
begins to form at an obtuse angle to the zygosporophore.
As development continues, the region of the zygosporophore
below what will become the collar of the zygospore swells
such that it is often twice the diameter (ca. 10 um) of
the supporting conjugant cell and the collar. A weak cir-
cumferential zone may be observed in the wall of the zygo-
sporophore where the collar will eventually detach. The
coiled appendage may be seen through the collar of the
attached, maturing zygospore (Fig. 5). After zygospore
release, it appears that the single appendage may not ex-
tend completely until stimulated by some mechanical
action. In contrast, the appendage of trichospores often
extends from the collar in its typical zigzag manner as
soon as the trichospore is released.
Acknowledgements: We are indebted to the National Science
Foundation for research grants GB-24947 and DEB-8019724
for support of our studies. The senior author acknow-
ledges the Research Services Council of Kearney State
College for partial support of this work. R. W. L. thanks
the Director of the Abisko Naturvetenskapliga Station for
use of laboratory facilities. Dr. Donald P. Rogers, Univ.
of Illinois, kindly provided the Latin for the diagnosis,
and we thank Dr. Leonard Ferrington, Kansas Biological
Survey, for identifying the type specimen host.
LITERATURE CITED
Kobayasi, Y., N. Kiratsuka, Y. Otani, K. Tubaki, S.
Udagawa, and M. Soneda. 1969. The second report on
the mycological flora of the Alaskan Arctic. Bull.
Nat. Sci. Mus. Tokyo, Japan. 12:311-430.
Williams, M. C., and Ri: W. Lichtwardt.. 1972. Infection of
Aedes aegypti larvae by axenic cultures of the fungal
genus Smittium (Trichomycetes). Amer. J. Bot. 59:
189-193.
MYCOTAXON
Vo. GX Wea NOhe lepp enlee ea Ls 2 October-December 1982
a
pe
MEGASPOROPORIA A NEW GENUS OF RESUPINATE POLYPORES
L. Ryvarden
Botanical, Institute, University of Osio,
P.O.Box 10945 Blindern, Oslo 3, Norwav
A ee or Wright and M. Raj chenberar
Departamento de Ciencias Riolégicas
Facultad de Ciencias Fxactas y “Naturales
University of Buenos Aires
Arcentina
SUMMARY
The genus Megasnorovoria Ryv. & Wriqht is cCescribed with
Poria setulosa Henn. as tyne species. The genus is charac-
terized by large nores and snores, clamned cenerative
hyphae and dextrinoid skeletal hynhae. The following new
combinations are proposed: Megasvoroporia cavernulosa
(Berk.) Ryv., Megqasporoporia hexaqonoides (Sneq.) Wright &
Rajch. and Megasporoporia setulosa (Henn.) Rajcn. Mecasvoro—
poria mexicana Ryv. is described as new.
Grammothele as defined by Ryvarden & Joharsen (1989:34-35)
included taxa with a resuvinate, voroid fruithody where
the basidia lined both the walls and the khottom of the
pores. Microscopically tne species were characterized by
dextrinoid skeletal hynhae and in many species also cendro-
hyphidia were present. The spores in all taxa were ellinp-
soid to cylindrical, non-amyloid and thinwalled with one
exception. Julich (1982) excluded G. macrospora Ryv. and
transferred it to Grammothelonsis dll. because of its
thickwalled, dextrinoid spores.
Among the remaining snecies G. delicatula (Hern.) Ryv. and
G. setulosa (Henn.) Ryv. are somewhat deviating as they
both have much larger pores and snores than the tyve
species of the genus, viz. G. lineata Berk. & Curt.
Further studies in the resuninate nolypores nave shown that
there are two more snecies which are closely related to
G. delicatula- and’ G. isetulosa. Weeteel that theses soe. =
constitute a natural taxon and propose a new genus to
accomodate them.
1) Work partially subsidized by a grant from the Consejo
Nacional de Investigaciones Cientificas y Técnicas.
2) Member of the "carrera del investigador cientifico" of
the above Consejo.
3) Fellow of the above Consejo.
es
MEGASPOROPORIA Ryvarden & Wright nov. gen.
Frutificatio resupinata, pori maqni, pori facies cremea,
cinerea ad pallidum cinnamomea, systema hynharum dimiticum
ad trimiticum, hyovhae generatoriae fibulatae, hyohae
skeletales crasse tunicatae, dextrinoiceae, hyphae
liganteae dextrinoideae, praesentes vel absentes, dendro-
hyphicia praesentia vel absentia, snorae magnae, hyalinae,
cylindricae, non-amyloiceae, tenuitunicatae.
Species typica: Poria setulosa Henn. Engl. Bot. Jahrb.
Coes LOD LS
Fruitbody resupinate, pores generally large, angular to
round, pore-surface cream, greyish to nale brown or
cinnamon, context usually very thin white to cream or
very pale brown. Hyphal system di-trimitic, generative
hyphae with clamvos, skeletal hynhae thickwalledc and
dextrinoid, branched vegetative hyphae which may be
interpreted as binding hyphae present in most svecies,
dextrinoid. Spores cylindrical, thinwalled and large,
non-amyloid and non-dextrinoid. Cystidia absent, dendro-
hvohidia present or absent, crystals usually present,
often abundantly in the subhymenium and the context.
On deciduous wood with a white rot. Predominantly a
tropical genus.
Remarks. The genus is characterized by its resupinate
fruitbodies, the large svores and pores besides strongly
dextrinoid skeletal hyphae. From Grammothele as defined
by its tyne svecies, it is senarated bv the larger
spores and pores and a more distincly voroid fruit-
body. In Grammothele the pores are in general small
to minute and very shallow, which give the hymenophore
a reticulate pattern or the aspect of low ridges more
than a distinctly poroid apnearance. The nresence of
dendrohyphidia in some of the species cCescribed here
seems to point to a relationship to Grammothele as such
organs in qeneral are rare among the volypores. Dendro-
hyphidia are often very difficult to observe in dried
polypores and they may have a wider distribution than
hitherto assumed. Fresh or properly dried specimens are
necessary to acertain their presence. Even if dendro-
hyphicia should be absent in some species we neverthe-
less feel that the characters mentioned above justify
the genus in its present circumscription.
The hyphal system is somewhat difficult to interpret.
The vegetative hynhae in some of the snecies show
transitions from unbranched long and thickwalled
skeletal hynhae to more sinuous hyphae with occasional
branching up to arboriform or irregularly hranched
hyphae. The latter, when observed in freefloating seq-
ments, may easily be taken as true bincing hyphae.
They should better be called branched skeletal hynhae
Since they often have in the lower part long unbranched
segments.
174
Key to species:
drei (POPCS 2-59 MM AW ELGG las 0ers 0s bre ete dle -B ta bieietal eo a DIR UIMNG i eaiie
i. Poresetessi Chane sani wide’ <.inKkssbene se eee eee
2. Hyaline, acute hyphal pegs abundant in the
VOresS seer cc ccc rece ccc cccccceseccecceed. M. Setulosa
2. Hyaline hyphal neqs absent or only very few present 3
3. Pore surface grey to greyish brown, pores 0,5-1] mm
wide, spores 16,5-22 um long, dendrohynhidia
ADS SN so: ave le oes buoys oie) 4,8 46 oleveeae eet ot Mame ka One Gar
3. Pore surface vale cream to ochraceous, vores 2-4
per mm, spvnores 10-16 (18) um long, dendrohynhidia
present along the pore edges ....1.M. cavernulosa
1. MEGASPOROPORIA CAVERNULOSA, (Berk.) Rvyv. Big secs
Comb.nov. - Basionym: Polyporus cavernulosus Berk.
Hook. J. Bots 822357 i856.) Portas delicartilasrtenns
Engl. Bot. Jahrb. 34:44, 1904. - Hexagonia heteropocra
Pat. J. Bot. (Morot) 3:166, 1889. = \Hexagonia bartlettisz
Mass. Bull, Misc. “Inform. Kew. 19082216, 1906%5= Poria
linearis Murr. Mycologia’ 12:303,. 1920.
Fruitbody annual, resupvinate, effused, adnate_and
coriaceous, up’ to 2 mm thick. Pore surface first white
to cream, then pale strawcoloured and finally when dry
and old, ochraceous to nale woody brown. Pores angular
anc shallow, 2-4 per mm, un to 1 mm deep, nore edcges
finely fimbriate in actively growing snecimens. Context
white to pale strawcoloured.
Hyphal system trimitic, generative hynhae thinwalled,
2-3 um-wide and with:-clamnps;9 often (citficult ,torting,
skeletal hyphae dominating, thickwalled, mostly
Sinuous and unbranched, 2-3 um wide and strongly dextri-
noid, binding hyphae or branched skeletal hynhae also
present, especially in the context, arboriform and 2-4
um wide, strongly dextrinoid. Dendrohynhidia present,
but difficult to observe in old and dry specimens, most
easily seen along the nore-ecges where they are abundart
in young and growing specimens, hynhodid to ventricose
with irregular branching in the upoer part; arising from
a clamp at the base, up to 25 um long. Spores. cylindrical,
hyaline, thinwalled and non-amyloid, variable in size,
10-16 (18) x 5-7 um, when young and immature apparently
somewhat ellipsoid; when mature loncer and slender.
Habitat: On deciduous wood. Distribution: Tropical
Africa and America.
Remarks. The size of the svores deserves some remarks.
The type of P. cavernulosus Berk. is not sterile as stated
ES
Fig.1. Megasporoporia cavernulosa A) section through the
hymenium. Coll. R. 10760, B) vegetative hyphae, C) spores.
From the type of P. cavernulosus, D-H) Spores, D) From the
type of Poria linearis, E) Andersson, 20,3,1980 (Ecuador),
F)Deighton 1.8.1949, (Sierra Leone), G) Ryvarden 10760 (Ken-
ya), H)From the type of Poria delicatula Henn.
LO
by Lowe (1966:130). After repeated examinations spores were
found that measured 9-12 x 4-5 um. The dextrinoid reaction
of the skeletal hyphae is very strong in this specimen. In
the type of Poria delicatula the spores measured 12-14 x
4-6 um while all intermediate sizes were found in others
specimens examinea. We believe that the medium sized
pores, the cream to ochraceous fruitbody and the strongly
dextrinoid skeletal hyphae are diaqnostic for this snecies.
The relatively large spore-variationr that can he ohserved
from the type of P. cavernulosus and of fresh snecimens
collected in Ecuador (see list below) probahly reflects
only different stages in development. After having examined
many specimens from Africa and America we came to the con-
clusion that a separation based on spore size was not vos-
sible.
The description given by Ryvarden & Johansen (1980:37) was
based partly on an undescribed species with almost the same
size of the spores but with yellow and non-dextrinoia
skeletal hyphae. This species will be treated in a later
paper.
Representative specimens: Brazil, Panuré, Ac ramos (tyne
of PP.) cavernulosus, K.); Bahia, Serra da. Acqua de Reda,
ZE7LU7 LOU e HLS? Virwin 2 (NY )-sCubas. (Col) SeWric hiteuge ou
(det. Ps cavernultosus by Berkeley) > Panama vy Marragante,
leg. H. Williams 3/1V/1908 (type of Poria linearis Murr.).
(NY) Bouador: Pastaza:, Curaray, 20/1I1/19S02 Mee)
Guyana: Tumatumari, beg. G. Linder, 16/1X%/1923. (NY).
Tanzania: Usambara, Bomole, May 1992, leg. G. Zimmermann
(evype: Of Portas delicatuia’ Henn. 7 (S$). Morogoro rploy.
UlLuguri Mts. Morning=Sice Res, Sta. 24/11/1973 7b. nyvar-
den 10944 (0). Zaire: ‘Shaba prov. Luisursiti, no date,
16q. 2Thoen=no® S43 17 (ER ;O)),
2. MEGASPOROPORIA HEXAGONOIDES (Speq.) Wright et
Rajchenberg comb. nov. Fig. 2.
Basionym: Poria hexagonoides Speg., An. Mus. Nac. Buenos
Aires*6 2 L7O=2 73935
Fruitbody annual, lignicolous, totally effused and attached
to substratum, circular to largely ellivsoie up sto 10 3
ecm, hard, coriaceous. Pore surface when fresh light ash
grey with lavender tints, when dry light greyish brown.
Margin absent or present and then cefined and regular,
always cream, up to 2 mm wide. Pores large, honey-combhed,
0>5=Tinmm wideseeontext, thin, Licht brown . 0,351, 7.10
thick. Tubes “unto. 2;5 mm long, concolorous wi thicontes.,
with triangular section. Alkali reaction negative.
Hyphal system trimitic. Generative hyphae clamped, branched,
thin-walled, 2,1-3,6 um diam.; skeletal hynhae branched or
not, with hyaline thickened walls or solic, 2,1-6,8 um
diam., dextrinoid; binding hyphae branched with long and/or
short branches, with hyaline thickened walls, 1,9-3,6 um
diam., dextrinoid, all the hyphal elements are arranged
intricately in the trama.
ids
section through the
A)
From the lectotype.
hexagonoides
G74 oC) ispores<
Megasporoporia
hypha
FLO.
hymenium, B)
L778
Hymenium 35-53 um thick. Basidia claviform 33,8-36,4 x
9,4-10,4 um, tetraspored, sterigmata triangular up to 6,2
um long. Basidiospores cylindric to slightly allantoid,
anpiculatec, hyaline, thin-walled, with or without oily
inclusions, 1¢,/6-21,6 * 5,2-6,8 um, inamyloid, incextri-—
noide, acyanonhilous. Basidioles cylindric to claviform
20,846, 90 Xe5,7 2-11, 4 um, with Gr withoub <istinct os.
drovs. Crystals polyedric, abundant, scattered between
hymenial elements and in subhynmenium. Cystidia ahsent.
Habitat: Prosopsis nigra and other unidentified anqicsverms
with a white rot. —- Distribution: Argentina: In suhxero-
phytic vegetation of the Chaco region (provinces of Salta,
Santa Fé and Tucuman).
Remarks. The spnecies has only slightly smaller spores and
pores than M. mexicana, but is easily identified in the
field because of its, ashy grey to lavender fruitbodies.
Lowe (1966:134) states that the isotype in BPI is sterile,
but in the holotyre in herb. LPS all hymenial elements
are present.
Representative specimens: Argentina, Salta, La Vina, leg.
Spegazzini, 1/1897 (HOLOTYPE LPS 25538), between Rosario
Ge la Frontera to G. Garmendia, leq. A. Okada VIII/1963
(BAFC 27916). Ibid., GUiemes, Mojotoro River, leg.
J. Déschamms SA=2910 5S/ITI/1976, on Living branches, of
Prosopis nigra (BAFC 27927). Santa Fé, Santa Margarita,
leg. G. Dominguez 2/X/1943 (BAFC 27918). Tucum&n, Dique
del Cadillal, .leq.. Ru Singer T=L566, 1O/VI/1oSAs (BARC
21919).
3. ‘MEGASPOROPORTA. MEXICANA Ryv. nov... Sp. Pigs 3.
Fructificatio resupinata, nori facies cremea, vori magni,
2-3 mm latus, Systema hypharum di ad trimiticum, hyvhae
generatoriae fibulatae, hynhae skeletales haud ramosae
vel ramosae et arboriformae, crasse tunicatae, dextrinoideae,
sporae hyalinae, cylindricae, non-amyloiceae 2%-26 x 6-9 um.
Typust Mexico, Vera’-Cruz, Coscontla, Municipio demituatuzcoa,
L6/V/1993, 2390 m ade. Coll. Ventura. 0370 .. Hototyoose.
herb. ENCB, isotyni in, herbaria 0, K anc, Bel.
Fruitbody annual, resupvinate, widely effused, in the type
ups towz0 cm long .ebo. cm wide sand 1 cm Enrich, toughened
coriaceous. Pore surface white to pale cream, pores
angular to round, 2-3 mm in diameter, tubes up to 2 cm
deep, cream coloured, with a few scattered low hyphal
pegs or warts in the upperpart. Context 2-490 um thick,
white ana fibrous, trama white and cense.
Hyphal. system di-(tri.2)-=mitic, -enerative, nyphaero.ins
walled, hyalire 2-4 um in wide and with clamms at the
septa, skeletal hynhae strongly dextrinoic, thickwalled
and of a variable shape, partly as long unbranched segments
arising from a clamn, up to 500 um long and with a rounded
apex, 3-7 um wide, partly as more sinuous seacments of
eo)
Fig. 3. Megasporoporia mexicana A) basidia, B) vegetati-
ve hyphae, C) spores. From the holotype.
180
variable length and with transitions to more narrow an@
dichotomously branched hyphae, 2-4 um wice. The latter type
of hyphae is interpreted more as upper encs of skeletal
hyphae more than true binding hyphae as the branching is
rather scarce. Basidia clavate, 35-45 x I0=13 um with 4
sterigmata, hyaline varaphysoic hyvhae present between the
basidia, 3-5 um wide. Spores cylindrical to allantoid, eine
walled non-amyloic, hyaline anc with smaller and larger
olldrops, 29-26 * 6-9 um. On decicucus wood. Known only
from the tyne.
Remarks. M. mexicana seems to be close both te M. setulosa
and M. hexagonoices. From the former it is separated ny
far less hyphal neqs, larger snores and pores. From the
latter it is separatec by larger vores anc a white to nale
cLean;r Colour, Of Ene fruitcody.
4, MEGASPOROPORIA SETULOSA (Henn.) Rajchenberq Fig. 4.
Nov. com. = Basionym: Portia setulose Menn.,- Ena. Bot.
Jahrb.) 28:321, 1901. — Trametes sussperpens Murrs Mycolegad
TPZ2¢1905 1920... involve fame. No Cescrin=ionm,
Fruitbody annual, lignicolous, totally effusecd, forming
small° cireular vatches finally coalescing “up to) 307 Ger,
corky. Pore surface white, cream, turning light brown upon
drying. Margin usually present, cGefinite,. regular, up to
1,7 mm wide, concolorous with hymenial surface. Pores
round, isodiametric, 1-2 per mm. Context thin, wo 7to oe
mm wide. Tubes up to 1 mm long. Alkali reacticn negative.
Hyvhal system dimitic, generative hynhae clamned, branched,
with thin hyaline walls, 1,6-3,1 um diam., and skeletal
hyphae slightly or not at all branched, 1,6-5)724um clam,
with hyaline thickened walls with lumina visible, or solid,
Gextrinoid. All elements are intricately arranged and
dissection is cifficulte. Hymenium 18,8-3255° um, thicz.
Baslidia: claviform, with or without. ofly irelusions, —c,
20,6 °X-6,0-9,4 “am, tetraspored, with straiont er curved
triangular sterigmata 7,8 x 2,6 um. Basidiospores cylind-
ric, apiculated, hyaline, with large cil Grovs7; thin wal tea,
L0=14,.0 x 4,2—5, 7 um, sinamyloid, acy.anonni lous’.
Basidioles claviform, with small or larqgqe.ocily inclusions,
V3,,5=31,2 x 5,2-10,4 um. Paraphyvsoids composec by generna=
tive hynhae that emerge between basicioles. Crystals noly-
edric, abundant, scattered in subhymenium and in trama,
Ji=-54 x lo-43 um. Hyohal pegs very abundant, Jeng and
rectangular 49-160 x 16-38 um easily seen under the lens.
Habitat. On angiosperms with a white rot. Distribution.
Pantropical.
Remarks. The snecies is usually recognized already in the
field because of the numerous and prominent hyphal pegs
inside the pores. The tyne of Poria setulosa was probably
lost in Berlin during the last world war, but trem the
descrivtion and the name there shoulée be no doubt that
Hennings name oricinally covered the taxon Cescribec here.
182
Representative specimens: Argentina: Missiones, Cataratas
del Iguazu, leg. M. Rajchenberg. 4/III/19890 (BAFC).
Corrientes: Mburucuya. Ea. Santa Teresa, leq. J. Wright,
Gomez & Del Busto. 28/1/1962 (BAFC). Venezuela: Bolivar
Halo de Vergarena. L. Gupps. /X/1954 (NY). Ghana: Ashanti
Region. Jimra Forest Reserve. 24/1IV/1974. Ryvarden 12719
(0). Kenya: Coast prove,Shimba Hills, Makadara Forest.
L4/LI/1973e Ryvarden 20234 (ol). Indias’ Tamiy Nadu,
Madurai distr. Tiger Shola. 17/VIII/197@. kK. Kolancdavelu
NGL 349 (0)5:
Acknowledgements
We wish to express our recognition to the Servicio Nacional
de Parques Nacionales, Argentina, for the facilities
afforded during our field trips in the Iguazu National
Park, and to the Consejo Nacional de Investigaciones
Cientificas y Técnicas for financial assistance and the
Fellowshiv granted to Mario Rajchenherq.
References
Julich, W. L9Sls Hagher taxa of Basidiomycetes. Bia,
My cols (65;
Lowe, J.L. 1966: Polyporaceae of North America. The genus
Poria, State Univ. Coll. Forestry Syrac. bie. Lecun.
Pub.) 690
Ryvarden, L. & Johansen, I. 1980: A preliminary polynore
flora ‘of (East, Arrica.-FPungiftiora’:
MYCOTAXON
Vou Vl NOS Lot pp mls 3=186 October-December 1982
SMITTIUM CELLASPORA, A NEW HARPELLALES
(TRICHOMYCETES) FROM A CHIRONOMID HINDGUT
MARVIN C. WILLIAMS
Department of Biology
Kearney State College, Kearney, NE 68847
This is the 19th recognized species of Smittium to be
described, and the 15th described from midge larvae
(Chironomidae, Ceratopogonidae). Previously described
species of Smittium from all dipteran hosts have been col-
lected from some 12 states of the U.S.A., including Alaska
and Hawaii, and from Greenland, Russia, Japan, and at
least five European countries.
Smittium cellaspora Williams, sp. nov.
Trichosporae ellipsoidales, (20-)29(-36) x (7-)8.5(-10)
um, appendiculo tenui pluries longitudinem sporae longo,
collare (5-)9(-10) x ca. 2.5 um. Thallus brevis, sparse
ramosus, paene omnibus cellulis fertilibus. Retinaculum
simplex, area adhesionis discformi. Zygosporae ignotae.
Chironomidarum larvalium incola.
Trichospores ellipsoidal (20-)29(-36) x (7-)8.5(-10) um
with a fine appendage several times the spore length and
which may exhibit a coiled appearance upon release of the
trichospore; collar (5-)9(-10) x ca. 2.5 um. Thallus
short, up to 300 um in length or possibly more, 6-10 um in
diam, sparsely branched, almost all cells fertile. Hold-
fast simple, tending to form a disklike adhesive struc-
ture. Zygospores unknown.
Holotype: Slide MIS-15-I deposited with R. W. Lichtwardt
at the University of Kansas, prepared from the hindgut of
a Chironomidae larva collected 7 February 1982 in the Sac
River, 1.0 km north of the Greene County line on highway
184
MO 13, Polk County, Missouri, USA.
Hosts: Few hosts were collected from the type site and
the one saved for identification is Sympotthastia sp. All
chironomid hosts from this site were taken from leaf packs
or algal growth attached to rocks in flowing water.
The spore size and unusual growth habit (Figs. 1-4)
distinguish S&S. cellaspora from any currently described
species. The only other species reported to have spores
in this size range is S. orthocladii Manier (1969)
{=Rubetella orthocladii Manier & Mathiez (1965) nom. nud.}.
However, S. orthocladii is reported to have thalli with
several basal branches growing in "bundles" which is dist-
inctly different from the present sparsely branched
species. Smittium orthocladii is described with tricho-
spores measuring 25-33.5 x 6-7.5 um with a collar (5-)8.7
(-10) um and is reported to produce an encrustment sur-
rounding the several basal branches. Smittium cellaspora
has a wider spore and produces a disklike holdfast pad
(Fig. 3) and no encrustment.
Smittium culicis has a similar spore shape; however,
the trichospore is smaller and this species normally is
found in mosquito larvae, while S. cellaspora is found in
chironomid larvae.
The simple holdfast of the new species tends to form a
disklike structure with an adhesive appearance as the
thallus matures (Fig. 3). The basal cell is 30 to 50 um
long and the other cells range from 20 to 45 um in length
and from 6 to 10 um in diam. The sparse branching pattern
is a characteristic of this species and each cell generally
is fertile or produces a branch which is fertile. Sporu-
lation occurs early in the growth pattern as shown by a
2-celled thallus each with a spore (Fig. 2). Such an
Figs. 1-4. Smittium cellaspora. 1. Trichospore with
spiraled appendage and characteristic collar, x1000. 2.
Thalli attached to chitinous hindgut lining, 225; note
2-celled thallus with 2 spores (arrow). 3. Branched
thallus showing the disklike adhesive holdfast, x675. 4.
Thalli illustrating usual growth pattern, note lower
thallus with each cell producing a spore, 225.
\
|
SS
i]
. |
Hi
186
abbreviated development is not generally reported in
Smittium spp., and early sporulation has been observed
only in laboratory infestations or axenic cultures. When
spores of another species, S. culisetae, were fed to Aedes
aegypti larvae in the laboratory, thalli were observed
rarely to sporulate at the 2-celled stage in the hindgut.
This precocious sporulation appears to occur when the fun-
gus is becoming established shortly before the host larva
is going to molt and is not the usual growth pattern.
Also, in axenic cultures of S. simlii, single-celled
germlings have been observed infrequently to produce a
spore with no further vegetative growth.
It is not uncommon to find more than one species of
Smittium growing in the same dipteran hindgut, therefore
it is possible that S. cellaspora could be present in guts
containing another Smittium sp. which produces more pro-
fuse growth. Under these conditions S. cellaspora may
be overlooked due to its limited growth form.
The photographs and measurements were made from living
material mounted in water. We noticed some shrinkage in
trichospore width (ca. up to 1 um) after the water mounts
were fixed in lactophenol-cotton blue.
Acknowledgements: I am indebted to the Kearney State
College Research Services Council, to the National Science
Foundation for grant DEB-8019724 (Supplement), and to
R. W. Lichtwardt for consultation and use of laboratory
space at the University of Kansas. I wish to thank
Dr. Leonard Ferrington for host identifications. Dr.
Donald P. Rogers, Univ. of Illinois, kindly provided the
Latin for the diagnosis.
LITERATURE CITED
Manier, J.-F. 1969. Trichomycétes de France. Ann. Sci.
Nat. ((Bov.9 sser:) 12, 102 565-672.
, and F. Mathiez. 1965. Deux Trichomycétes
Harpellales Génistellacées, Parasites de Larves de
Chivonomides. “Ann: ‘Sci: “Nat; “(Bot. Ser. 12.
6: 183-196.
|
MYCOTAXON
ViOdEe AV Le NOwiel “pp. 187-191 October-December 1982
NOTES ON HYPHOMYCETES. XLII.
NEW SPECIES OF ACRODICTYS AND PSEUDOSPIROPES FROM
SOUTH AFRICA.
G. Morgan-—Jones
Department of Botany, Plant Pathology and Microbiology,
Auburn University Agricultural Experiment Station,
Auburn University, Alabama 36849, U.S.A.
ABSTRACT
Acrodtetys etekerti Morgan-Jones and Pseudospiropes falcatus
Morgan—Jones, two new species, are described and illustrated from
collections made on decorticated twigs and wood in South Africa.
INTRODUCTION
In earlier papers in this series Morgan-Jones and Sinclair
(1980a, 1980b) described new dematiaceous hyphomycete taxa, belonging
to the genera Stachybotrys Corda and Custingophora Stolk, Hennebert
and Klopotek respectively, from collections made on decorticated
wood in South Africa. Continued exploration of this substrate has
yielded collections of further undescribed species of which two,
belonging to the genera Acrodictys M. B. Ellis and Pseudospiropes
M.D. Ehlisseare descripeds herein.
TAXONOMIC PART
Acrodtetys etekerit sp. nov. (Fig. 1).
Coloniae effusae, atrobrunneae vel atrae. Mycelium partim super-
ficiale, partim immersum, ex hyphis ramosis, septatis, pallide
brunneis, levibus, 2 - 3um crassis compositum. Conidiophora macrone-
mata, singula vel 2-3 fasciculata, non ramosa, ex lateribus hypharum
oriunda, recta vel flexuosa, erecta, brunnea vel atrobrunnea, laevia,
continua vel septata, usque ed 40m longa, 5 - 6ym crassa, cum 0 - 2
proliferationibus terminalibus successivis. Cellulae conidiogenae
monoblasticae, in conidiophoris incorporatae, terminales, determinatae
vel percurrentes. Conidia solitaria, in apice conidiophori oriunda,
Sicca, subglobosa vel late pyriformia vel turbinata, muriformia,
levia, brunnea vel atrobrunnae, 38 - 64m longa, 22 - 42um crassa,
Alabama Agricultural Experiment Station Journal Series No. 6-82257
188
————
10pm
FIGURE 1. Acrodtetys etckerit
appendicibus 2 - 7, pallidebrunneis, 7 - 16um longis, 2 - 3um
crassis; ad basim truncata.
In ramulis emortuis decorticatis, Debengeni Forest Reserve,
Magaebaskloof, N. E. Transvaal, South Africa, August 17, 1979, R. C.
Sinclair, AUA, holotypus.
Colonies effuse, dark brown to black. Mycelium partly super-
ficial, partly immersed in the substratum, composed of branched,
septate, pale brown, smooth, 2 - 3yum wide hyphae. Conidiophores
macronematous, mononematous, single or in fascicles of two or,
rarely, three, unbranched, arising laterally from the hyphae, straight
or flexuous, erect, brown or dark brown, smooth, non-septate or
septate, up to 40m long, 5 - 6ym wide, with occasionally up to two
terminal successive proliferations. Conidiogenous cells monoblastic,
integrated, terminal, determinate, or proliferating percurrently.
Conidia solitary, formed singly at the apex of each conidiophore,
dry, subglobose or broadly pyriform or somewhat turbinate, muriform,
smooth, brown to dark brown, 38 - 64ym long, 22 - 42um wide, bearing
2 - 7 pale brown appendages distally and laterally, 7 - 16um long,
2 — 3um wide; base truncate.
On decorticated twigs; South Africa.
Collection examined: Debengeni Forest Reserve, Magaebaskloof,
No EB. Transvaal, South Africa, August 17, 1979, R. C. Sinclair, AUA,
holotype.
Acrodtetys etckerit is named in honor of Professor Albert Eicker,
Department of Botany, University of Pretoria, through whose good
offices the collections of hyphomycetes on which these studies are
based were made.
Among species of Acrodictys, A. etekerti resembles both A.
brevtcornuta M. B. Ellis and A. appendtculata M. B. Ellis in possess-—
ing a number of conidial appendages. It is morphologically quite
similar to A. appendtculata in particular but differs from it in
having much larger conidia, usually more numerous conidial appendages
distributed both laterally and distally, and possessing percurrently
proliferating conidiophores. In the latter characteristic it re-
sembles several non-appendaged species of Acrodietys. In shape
of conidia and distribution of appendages A. etekerii bears a broad
similarity to Piricauda ecochinenis (Subram.) M. B. Ellis.
Pseudosptropes falcatus sp. nov. (Fig. 2).
Coloniae effusae, olivaceo-brunneae vel fuscae, velutinae vel
pilosae. Mycelium immersum, ex hyphis ramosis, septatis, pallide
brunneis, levibus, 2 - 3.5um crassis compositum. Conidiophora
macronemata, mononemata, erecta, recta vel leniter flexuosa, crasse
tunicata, septata, brunnea, apicem versus pallidiora, laevia,
cicatricibus conidialibus praedita, usque ad 190um longa, 4 - 6um
erassa, basi unterdum ad 8yum inflata. Cellulae conidiogenae
LOO
Pseudosptropes faleatus
FIGURE 2.
192
polyblasticae, in conidiophoris incorporatae, terminales, sympodiales,
eylindricae, usque ad. / cicatricles. Conidia solitaria, sicca,
acropleurogena, simplicia, cylindrica vel fusiformia vel falcata,
basi truncata, pallide brunnea, laevia, 5 - 7 septata, 32-41 x4 -
Sum.
In ligno emortuo, Pienaar's River, Pretoria, South Africa, August
IS, tose Cc. oineleiys AUAT hoLoLypus.
Colonies effuse, olivaceous brown to dark blackish brown, velvety
or hairy. Mycelium immersed, composed of branched, septate, pale
brown, smooth, 2 -— 3.5yum wide hyphae. Conidiophores macronematous,
monomematous, simple, erect, straight or slightly flexuous, thick-
walled, septate, brown to pale brown, paler towards the apex, up to
190um long, 4 - 6ym wide, up to 8ym wide at the swollen base. Conidio-
genous cells polyblastic, integrated, terminal, sympodial, cylin-
drical, bearing a number of thin, flat, small, dark, very slightly
protruding scars. Conidia solitary, dry, acropleurogenous, simple,
cylindrical to fusiform, most frequently falcate, base narrowly
bruncate, pale brown, smooth, 5 - 7 septate, often bearing a small,
highly refractive spot at the extreme apex, 32 - 41 X 4 - Sum.
On dead wood; South Africa.
Collection examined: Pienaar's River bank, junction of Bronk-
Forevoprus.. head. Pretoria, south Africa, Ausust. 13, 1979, R. Cc.
Sinclair, AUA, holotype.
Pseudosptropes falcatus is easily distinuishable from other
species of Pseudosptropes possessing relatively thin-walled,
septate conidia, as opposed to the thick-walled, pseudoseptate
conidia of P. nodosus (Wallr.) M. B. Ellis and P. stmplex (Kunze)
M. B. Ellis, by the leneth and shape of its conidia. It most closely
resembles P. rousselianus (Mont.) M. B. Ellis, which also, inciden-
tally, has often a small refractive spot at the apex of its conidia.
ACKNOWLEDGMENT
I thank my former graduate student, Mr. Robert C. Sinclair,
for the opportunity to examine collections made by him in South
Africa. Dr. J. Leland Crane kindly reviewed the manuscript.
REFERENCES
MORGAN-JONES, G. and R. C. SINCLAIR. 1980a. Notes on Hyphomycetes.
XXXTII. Stachybotrys sphaerospora sp. nov. from South Africa.
Mycotaxon 10: 372-374.
MORGAN-JONES, G. and R. C. SINCLAIR. 1980b. Notes on Hyphomycetes.
XXXVI. A new species of Custingophora. Mycotaxon 11: 443-445.
MYCOTAXON
VOTERS eNO 19 291-96 October-December 1982
eee nee
NOTES ON HYPHOMYCETES. XLIII.
CONCERNING CHAETOPSINA ROMANTICA.
G. Morgan-—Jones
Department of Botany, Plant Pathology and Microbiology,
Auburn University Agricultural Experiment Station,
Auburn University, Alabama 36849.
ABSTRACT
Chaetopsina romantica Rambelli and Lunghini is described and
illustrated from a collection on leaves of Magnolta virgintana L. in
Alabama. The collection represents the second record of the fungus
and the first from North America.
INTRODUCTION
The genus Chaetopsitna Rambelli, which remained monotypic for well
over a decade after its establishment in 1965, has had a number of
species added to it during the 1970's (Matsushima, 1971; Rambelli and
Lunghini, 1976, 1979; Sutton and Hodges, 1976; Morgan-Jones 1979).
When described, each species was known from but a single collection
and this still remains the case with the majority. The exceptions
are the type species, C. fulva Rambelli, which is now known from dead
leaves in Italy (Rambelli, 1956), Papua-New Guinea (Matsushima, 1971),
U.S.A. (Pirozynski and Hodges, 1973), Japan (Matsushima, 1975),
Taiwan (Matsushima, 1980) and from soil in Canada (Barron, 1968), and
C. ramtfera Matsushima, which is known from dead leaves in Papua-New
Guinea (Matsushima, 1971), Brazil (Sutton and Hodges, 1976), Taiwan
(Matsushima, 1980) and from dead wood in the Ivory Coast (Rambelli
and Lunghini, 1979). We have no knowledge of the geographical dis-
tribution of the other taxa.
Chaetopsina romantica Rambelli and Lunghini was described from
unidentified bark fragments of forest floor litter collected in the
Tai National Forest, Ivory Coast, Africa, in 1976. It has not, to my
knowledge, been reported elsewhere to date. I have, however, en-
countered the fungus during an investigation of the microfungi occurr-
ing on leaf litter of sweetbay (Magnolta virginiana L.) in Alabama.
A comparison of the Alabama material with the description provided
by Rambelli and Lunghini (1979) indicates a number of important
Alabama Agricultural Experiment Station Journal Series No. 6-82259
oS
characteristics of the conidiogenous cells not documented by these
authors as well as a number of morphological features and vari-
abilities. A new description and illustration of the fungus is there-
fore presented here.
TAXONOMIC PART
Chaetopstna romanttea Rambelli and Lunghini, Trans. Br. mycol. Soc.
(2: 391, LOT9 (Rie. A).
Colonies effuse, hairy, glistening, brown to dark brown. Mycelium
partly superficial but most immersed, composed of septate, branched,
very pale brown, smooth, 2 - 3.5um wide hyphae. Setae formed directly
from the mycelium or from a cluster of a few to many swollen, subglobose,
yellowish to mid brown, thick-walled, 4 - 6ym wide cells, scattered,
solitarysor sometimes in pairs, erect’, mostly straight, verruculose
distally, thick-walled, brown, tapering gradually towards the apex,
bearing up to 16 septa, septae at closer intervals near the apex, up
to 370um long, 8 - 10um wide in the middle part, up to l4um wide at
the slightly bulbous base. Conidiogenous hyphal elements arising
laterally towards the middle of the setae or at various levels in the
distal portion, or terminally, simple or branched, subhyaline to very
pale yellowish, cylindrical, smooth-walled, septate, of varying
teneth. Conidiogenocus cells mono or polyphialidic, discrete, deter—
minate or nondeterminate, narrowly ampulliform or frequently assuming
an hourglass-shaped configuration, borne terminally or laterally on
the conidiogenous hyphae or, occasionally, directly from the setae,
bearing a discernible but Non-flaring collarette, 7 - 18 X 2 - 2.5um.
Sometimes integrated, intercalary cells of the conidiogenous hyphae
ean be fertile. Conidia enteroblastic, hyaline, aseptate, straight,
smooth, Cylindrical, obtuse av cach end, jf = 9 X 1.5 = 2um:
On bark and leaves; Africa and North America.
Collection examined: on dead leaves of Magnolta virgintana L.,
off Rt. 50, 5ml south of Lafayette, Chambers County, Alabama, July
26, 1979, G. Morgan-—Jones, AUA.
The binomial Chaetopsina romantica is being used for the Alabama
collection advisedly. Although the conidia are identical the conidio-
genous apparatus differs substantially from that in the type de-
scription. Of particular note is the sometimes indeterminate nature
of the conidiogenous cells which, as a result of sympodial extension,
increase in length, produce several conidiogenous loci and thereby
become polyphialidic. The peculiar hourglass shape of some conidio-
genous cells is evident in the illustration provided by Rambelli
and Lunghini (1979) but no mention of this is made in their de-
scription. This type of configuration is usually evidence, for per-
current proliferation of phialides but no discontinuity in the
periclinal wall at the point of constriction is discernible under
cne-resoulnvion limits of ‘the light microscope. in this instance.
The presence of conidiogenous cells that are polyphialidic in nature,
as well as fertile intercalary cells in the conidiogenous hyphal
194
Chaetopstna romantica
FIGURE 1.
i35
elements, in the Alabama material may reflect especially favorable
envirommental conditions rather than any fundamental genotypic
variance. This does, however, raise a question concerning the dis-
tinctiveness of Chaetopstna. As Rambelli and Lunghini (1976) have
pointed out the possession of polyphialides by Chaetopstis Greville
was regarded as the main differentiating characteristic between
the two genera. Chaetopstna tvortensts Rambelli and Lunghini also
possesses polyphialides [Rambelli and Lunghini (1976) illustrate
such structures although in their type description the conidiogenous
cells are described as being monophialidic]. I should also note in
passing that it appears that the phialides of C. tvortensts can
proliferate percurrently (there is, in fact, much similarity between
the conidigenous apparatus of C. tvoriensts and that of the Alabama
collection described herein but the conidia of that species are much
smaller). In both Chaetopsts grisea (Ehren.) Sacc. the type species
of Chaetopsts, and Chaetopsina auburnensts Morgan-Jones conidiogenous
cells are sometimes replaced by setose lateral branches. The primary
distinction between these two genera lies in the morphology of the
lateral, phialide-bearing branches; in Chaetopsts they are acutely
divergent, robust, and thick-walled towards their point of origin
whereas in Chaetopsina they are, generally, flexuous and thin-walled,
usually orientated more or less parallel to the seta, particularly
when borne towards the middle.
ACKNOWLEDGMENT
I thank Dr. Carol A. Shearer, University of Illinois, for review—
ing the manuscript.
REFERENCES
BARRON, G. L. 1968. The genera of Hyphomycetes from soil. Williams
and Wilkins, Baltimore.
MATSUSHIMA, T. 1971.Microfungi of the Solomon Islands and Papua-—New
Guinea. Kobe. Published by the author.
MATSUSHIMA, T. 1975. Icones microfungorum a Matsushima lectorum.
Kobe. Published by the author.
MATSUSHIMA, T. 1980. Saprophytic fungi from Taiwan. I. Hyphomycetes.
Matsushima Mycological Memoirs 1.
MORGAN-JONES, G. 1979. Notes on Hyphomycetes. XXXI. Chaetopsina
auburnensis sp. nov. Mycotaxon 8: 411-416.
PIROZYNSKI, K. A. and C. S. Hodges. 1973. New hyphomycetes from
Soura Carodine. Can. Jy Bot. SL: 157-173.
RAMBELLI, A. 1956. Chaetopsina nuovo genere de Ifali Demaziacei.
Atti Accademia Scienze dell ‘Istituto di Bologna 15: 1-6.
196
1976. Chaetopstina ivortensts a new
RAMBELLI, A. and D. LUNGHINI.
Morr. pot. weal. 210:
species of Dematiaceous Hyphomycete.
2532250
RAMBELLIT, A. and D. LUNGHINI. 1979. Chaetopsina species from
tropical forest litter. ‘Trans. Br. mycol. soc. 2. 4oi-494.
1976. Eucalyptus microfungl:
SUTTON, B. C. and CAS. HODGES.
Nova Hedwigia 27:
some setose Hyphomycetes with phialides.
343-352.
MYCOTAXON
VO Vie NOwils pp. l97-23d October-December 1982
ee ee ee
PHYCOSYMBIODEMES IN PSEUDOCYPHELLARIA IN
NEW ZEALAND
1 7
x x
B.RENNER and D.J.GALLOWAY
*1 Fachbereich Biologie der Universitat Marburg
Lahnberge, D-3550 Marburg, Germany (Federal)
*2 C/- Botany Department, British Museum (Natural
History), Cromwell Road, London SW7 5BD
England
SUMMARY
Three instances of joined thalli in Pseudocyphellaria
from New Zealnd are presented. Details of developmen-
tal morphology and results of phytochemical investi-
gations are given and discussed in connection with
recent findings in other genera. The importance of
chemical characters especially, in evaluating influen-
ces of different symbiotic algae is underlined.
Phycosymbiodeme is proposed as a functionally descrip-
tive term for joined thalli occuring most commonly in
the suborder Peltigerineae and especially in the fa-
milies Peltigeraceae and Lobariaceae.
The new species P.allanii D.Galloway, P.margaretae
D.Galloway, and P.murrayii D.Galloway, are described
and the new combination P.rufovirescens (Church. Bab.)
D.Galloway, proposed.
INTRODUCTION
Recent discussions of morphotypes in Stteta (James
and Henssen 1976), and in Pelttgera (Brodo and
Richardson 1979) expanding earlier accounts of
joined thalli in Sttcta (Wilson 1891, Dughi 1944)
and in Lobarta (Dughi 1937, Jordan 1972)have made
necessary a re-evaluation of the problem of inter-
action between mycobionts and phycobionts in the
lichen symbiosis. Detailed comparative studies on
198
composite thalli in the order Peltigerineae, and
in the family Pannartaceae (James and Henssen 1975,
1976) showed that both anatomy and morphology of
certain taxa in these groups may be influenced by
the phycobiont present in the symbiotic system.
Different genera of phycobiont exert distinctive
morphogenetic pressures (e.g. the development of
stalked fronds) on the same mycobiont, thereby
stimulating the production of joined thalli under
certain, usually favourable, ecological conditions.
Studies of Brodo and Richardson (1979) on the de-
velopment of subfoliose cephalodia in Pelttgera
aphthosa and Coccomyxa-containing lobes growing
from them, support the notion of differing lichen
morphologies resulting from a particular mycobiont
associating with different phycobiont genera in a
single autonomous plant. Further they discuss a
deviating secondary metabolism as a possible cause
of two different symbiotic unions, a speculation
having important taxonomic implications.
In Pseudocyphellarta Vainio, in New Zealand (Gallo-
way in prep.) several species pairs exist, compri-
Sing a fertile (parent) species, and a correspon-
ding asexual (derived) species possessing either
isidia or soredia but usually rarely or never fer-
tile. Examples of usually such species pairs are
P.homoeophytta (fertile)-P.deltsea (isidiate) and
P.faveotata (fertile)-P.granutata (sorediate). It
is thought that the asexual species (often more
widely distributed) is derived from the sexual
species, and from an evolutionary point of view
is regarded as having arisen at some time subse-
quent to the emergence of the fertile parent spe-
cies. Interestingly enough, in the case of P.de-
Ztsea which is widespread in New Zealand, the sub-
antarctic islands, south-eastern Australia and in
southern South America, the fertile parent species
P.homoeophyttlta, is known only from New Zealand.
Besides conventional species pairs in Pseudocy-
phellarta, field studies disclosed also the exis-
tence of joined thalli composed of closely simi-
lar species apparently differing mainly in the
presence of a green or blue-green phycobiont. Sub-
sequently, free-living individuals of the blue-
green phycobiont-containing species were dis-
199
covered and are described in this paper since the
green phycobiont-containing species in all cases
was already named.
In at least five instances joined thalli are know
and doubtless more will be found in New Zealand
and elsewhere. In this paper observations on three
of these are presented together with details of
their chemistry , morphology and taxonomy.
James and Henssen (1976 pp 53-56) record evidence
Giaas@aoqntsand humidity giving rise \to morphotypic
Variation in species of Pseudocyphellaria (P.
punetulata and P.junghuhntana) in south-east Asia,
with P.punetulata (green phycobiont) occuring in
high light situations in canopy branches and P.
junghuhntana (blue-green phycobiont) growing in
low-light, high humuidity sites at the base of
the tree or on adjacent rocks. Between the two
extremes of habitat and microclimate there exist
a series of intermediates which "....freely devel-
Op pycnidia and ascocarps and, significantly, there
is relatively little alteration in their respec-
tive thalline anatomy and morphology according to
which of the algal partners predominates ...."
( James and Henssen p.55). The situation with
joined thalli found in New Zealand is somewhat
different since both free-living green-phycobiont,
and blue-green-phycobiont species can exist in
similar habitats alongside examples of joined
thalli, and there doesn't seem to be any particular
microclimate factor potentiating the formation of
joined thalli. However it must be noted that the
joined thalli so far known are all from moist,
evenly humid, habitats in moderate shade either
close to sheltered streams, or in + deep gorges
or in areas of wet forest on slopes often shron-
ded in cloud. Since both green, and blue-green
phycobiont-containing components of joined thalli
are capable of an independent existence and have
certain characters which allow their taxonomic
separation they are given, for the time being, se-
parate species names. The taxonomic status of the
7Oined thalli is still a matter for major discus-
sion and no single name can be ascribed to it un-
der the existing rules of Botanical Nomenclature.
In our treatment we recognise the names of the
200
two components (which appear to differ mainly in
terms of the phycobiont present) within quotation
marks, Since it cannot be proved unequivocally
(although it is strongly suspected) at present
that the same fungus is common to the two compo-
nents of the joined thalli. It is quite possible
that already well known green phycobiont-,or blue-
green phycobiont-containing species will be found
joined to similar species differing in the nature
of the phycobiont. Such a phenomenon seems to
occur with some frequency in the families Peltt-
geraceae and Lobartaceae, especially in areas of
rich speciation where it appears that species are
in the process of emerging to exploit a particu-
lar microhabitat and/or microclimate.
TERMINOLOGY
The use of chtmera (James 1975) and chtmerotd
gssoctatton “(Brodo and) Richardson 119:79)> issmis—
leading and best avoided. Winkler (1907) first
introduced the term to describe plants composed
of two or more idiotypes derived from somatic
mutation, segregation or grafting. Such a defini=
tion cannot be applied to any known lichen thal-
lus. On the other hand the term morphotype pro-
posed by James and Henssen (1976) to emphasize
the dimorphic character of joined thalli takes
into consideration only one aspect of these com-
posite organisms, although admittedly according
to our present knowledge, the most striking as-
pect. We here propose an alternative term phyco-
symbtodeme to encompass all the peculiarities of
joined thalli presently known. Swinscow (1977)
used the term phycotype which is close to the term
proposed here, but does not fully agree with our
definition. The use here of deme rather than type
as an ending in the terminology proposed above
frees the term from any taxonomic or systematic
connotation which may inadvertently be placed on
it. Deme is a suitably neutral suffix which indi-
cates a specific type of relationship between a
group, of individuals of a specifie taxon,,e-9*%
chemodeme, morphodeme etc.. It deserves a wider
use in descriptive lichenology.
201
Phycosymbtodeme. In a number of lichens especially
in the Pelttgeraceae ,and Lobartaceae the particu-
lar morphological expression of the symbiosis is
causally related to the species of symbiotic al-
gae. Natural phycosymbiodemes always occur in
pairs and are distinct products of lichenisation
of one mycobiont with two different algae, one
belonging to the Cyanophyceae the other to the
Chlorophyceae. Any dissimilarity may be caused by
physiological and/or morphological and/or anato-
mical and/or cytological differences between the
two lichenised states. If one of the two phyco-
symbiodemes of a pair contains green algae, and
the other blue-green algae then the first one is
called a chlorosymbiodeme, the second one a cy-
anosymbiodeme. A prtort the quantity of distin-
guishable algal species in either phycosymbiodeme
tsenoc-lamited ;
MATERIAL AND METHODS
MATERIAL
Living material of joined thalli were obtained
from several localities in New Zealand and were
sectioned within 4-6 weeks of collection. Mate-
rial of the pair "P.margaretae/P.pubescens" came
from Leptospermum bark, Black Hill, Nelson Lakes
National Park (South Island) in 1978, material
ODM Mur rautT/ e.ruyouLtrescene \sLromitwigs sor
Lophomyrtus bullata, Mangaotaki River, King Coun-
try mtINOreh iisland) in 1978, and imaterial of
PPRallonte/Pecortaeca . Erom bark of Myrsine au-
stralts, Peel Forest, Little Mt Peel (South Is-
Langd)win e977.
METHODS
Morphology:Freezing-microtome sections were
mounted in lactophenol/cotton-blue. Habitat pho-
tographs were taken with a Wild M/ microscope,
photographs of micromorphology with a Wild M20
microscope. Scanning electron microscopy of air-
dried thallus fragments was performed with an
AMR 1200 SEM (Fa. Leitz). Lichen material was
attached to stubs without prior chemical fixgtion
and coated with a thin gold layer (c.6 x 10 mm
thick)using a sputter coater with gold target as
cathode (sputter coater Fa. Balzer, glow dis-
202
charge cin, aimat 10505 mbarrands1e0svolcs):
Chemistry: Secondary metabolites were analysed
by TLC in solvent systems A,B and C (Culberson
1972). Identification of some lichen compounds
by quantitative isolation from thin-layer chro-
IMatograms followed the method of Renner and Ger-
stner (1978). UV/VIS-absorptionspectra were re-
corded on a DMR 10 spectrophotometer (Fa. Zeiss)
and mass spectra on a Varian CH7 . masspectrome-
ter (using direct inlet system and PFK as inter-
nal standard; energy of the ion beam was 70 eV,
ion acceleration voltage was 3 kV).
Trimethylamine was detected by gas chromatogra-
phy on a Varian Aerograph 1400, equipped with a
flame ionistaion detector (column: Poropak T;
Carliierudas Na eo. umn Tenperacire. 50°C; injec=
tion temperature: 240°C; range: 2 x 10 [amps/
TV] ie
TAXONOMIC PART
Pseudocyphellarta allantt D.Galloway sp.nov.
Species corticola Novae Zelandiae Pseudocy-
pheltartam cortaceam (J.DoHook. & Taylor) DeGal-
loway & P.James, simulans sed pagina superiori
plumbea,. scabrida, phycobiontis coerulescenti-
bus Nostocaceis differt.
Holotype: New Zealand. Canterbury, Mt Peel, on
track to Emily Falls, Peel Forest. On fallen
Pseudowtntera on bank of stream, 20 March, 1979.
D.J. Galloway, CHR 343256.
Thallus orbicular to spreading, + entangled, to 18 cm
diam., loosely to closely attached, corticolous. Lobes
linear-clongate (3=)5-8" (14) mm wide; 1.5" = CG _cmalong7s
subcanaliculate, subdichotomously branching, discrete,
margins entire, sinuous, slightly thickened below, faint-
ly white-pubescent towards apices. Upper surface dark
greyish-blue to blue-green, tinged brownish at margins
when wet, pale fawnish-grey when dry, minutely scabrid,
coriaceous, uneven or very Slightly wrinkled-undulate,
without isidia, soredia or pseudocyphellae. Medulla white.
Phycobiont blue-green, Nostoc. Lower surface densely to-
mentose to margins, occasionally glabrous centrally, pale
203
buff at margins, dark brown to black centrally, tomentum
thick, silky, pale whitish to dark brown, occasionally in
scattered, squarrose tufts. Pseudocyphellae white, scatter-
ed, conspicuous, round to irregular, 0.5 - 2.5 mm wide,
plane to concave with a rised margin at maturity, often
sunk in tomentum.
Apothecia not seen. Pycnidia sparse to frequent, immer-
sed, 1.5 mm diam., visible as hemispherical swellings on
the lower surface.
Chemistry: 7B-acetoxyhopane-22-o0l and hopane-15a-22-diol.
Specimens examined: New Zealand. South Island. Canterbury:
Peel Forest, D.J.Galloway (CHR)
P.allanit is very closely related to P.cortacea
and often grows attached to this species. It has
so far been found only at Peel Forest on the
eastern slopes of Mt Peel. Here it most commonly
occurs as an epiphyte of Myrsine australis and
when wet is readily distinguished from P.coriacea.
It associates with P.epitsttcta and P.homoeophyl-
t@but its ecology is still poorly known. It is
DenearLOrDre Heh Allan. Dirst Director :of Botany
Division, >. ook, who tin the 1920's eescribed
the vegetation of Mt Peel, and who later promo-
ted interest in New Zealand lichens.
Pseudocyphellarta margaretae D.Galloway sp.nov.
Species corticola Novae Zelandiae Pseudocy-
phellartam pubescens (MU11. Arg.) D.Galloway & P.
James, simulans, sed lobis imbricatis, rotunda-
tis, pagina superiori plumbea, non faveolata,
tomento sericea dense obtecta, phycobiontis coe-
rulescentibus Nostocaceis differt.
Holotype: New Zealand. Nelson. Lake Rotoiti, Nel-
son Lake National Park, Peninsula Walk, on bark
Of Leptospermum erteotdes, 29 February, 1980. D.
ce neal lowady ., CHR 343279 ,°\lsotype in BM.
Thadl us orbicular, rosette-forming to spreading, 6-10
(-16) cm wide, loosely attached, corticolous. Lobes short,
rounded, 0.3 - 0.8 mm wide, imbricate or contiguous, mar-
gins entire, sinuous, shallowly incised, often subascen-
dent, densely white tomentose. Upper surface dark livid
leaden-grey or greyish-brown when wet, pale brownish-grey
204
or pinkish-fawn when dry, + uniformly tomentose, tomentum
long, white, silky, cortex below tomentum smooth, undulate,
uneven, shining, never scabrid. Medulla white. Phycobiont
blue-green, Nostoc. Lower surface pale brownish or whitish
at margins, dark brown centrally, irregularly ridged or
wrinkled-faveolate towards margins, + uniformly tomentose,
tomentum thick, whitish, silky, uneven, often developing
in long, tufted fascicles towards margins. Pseudocyphellae
yellow, tiny, 0.1 -0.3 mm diam., scattered or frequent,
sunk in tomentum.
Apothecia not seen.
Chemistry: tenuiorin, methyl gyrophorate(+), methyl lecano-
rate, methyl evernate, pulvinic dilactone, pulvinic acid,
calycin, nOrstLleticy) constictic, Ccryptostictic and cticere
acids, hopane-15a-22-diol, hopane-6a,78,22-triol.
Specimens examined: New Zealand. North Island. Hawkes Bay:
Kuripapango, J.K.Bartlett (BM). South Island. Nelson: Cobb
Valley, J.K.Bartlett (CHR 266031), Cobb Dam, J.K.Bartlett
(CHR 266029), Lake Rotoiti, West Bay, D.J.Galloway (CHR
343216), St Arnaud, Black Hill, D.J.Galloway (CHR 343217,
343274), Tophouse, W.Martin (CHR 375474).
P.margaretae is a very characteristic species
apparently most common in + ‘subalpine habitats ain
north-west Nelson, New Zealand. It is an eviphyte
of Leptospermum ertcotdes, L.scopartum and Olea-
YPLOsaVLCennafol71dgsan, habitats) modiiiedsby sister.
It is tolerant of moderate shade and is found in
areas of Aigh rainfall and, humidity. Although
often reaching a considerable size it is nota
common lichen in the habitats in which it grows.
It associates with the following lichens: Anzvta
jamestt,, Nephvoma australe, Nswcellulosum, Niltepe-
dophyllum,, Pannoparmelta .angustara.: P.welsonr,
Pseudocyphetlaria ecarpoloma, P.colensot, P.corce-
mata, Pecrocgta, P.granulata;, neglecta, New
bescens, P.nubella, Psoroma ieuphy lum, ih. adupvreteez
and, Papa ti iaum.
It is related to P.pubescens and the two species
have been found attached to each other. However
it differs from P.pubescens in the following Te>=
spects: the lobes are shorter, and more rounded-
imbricate, the upper surface is not visibly fa-
veolate or even markedly ridged, the surface of
the lobes is uniformly tomentose and the colour
205
of the upper surface when wet is a livid, leaden-
grey-brown, because of the presence of the blue-
qreen-phycobiont. In contrast, P.pubescene is
conspicuously green when wet, has longer and
narrower lobes which are faveolate-ridged, the
upper surface is often glabrous and the cortex. +
GiuStIinGtL ly scabrid. Lt \is Often tertile, and the
marginal apothecia have black discs usually cover-
ed with a grey-white pruina. P.margaretae has not
yet been found fertile.
It is named for Mrs Margaret Bulfin (née M.J.A.
Simpson) who has worked for many years on the ve-
getation and flora of the Nelson Lakes National
Pack’.
Pseudocyphellarta murraytt D.Galloway sp. nov.
Species corticola Novae Zelandiae Pseudocy-
phellartam rufovtrescens (Church.Bab.) D.Galloway*,
simulans sed pagina superiori plumbeus vel cine-
rascens, undulata vel subfaveolata, maculata,
phycobiontis coerulescentibus, Nostocaceis, sub-
tus tomento sericea dense obtect, differt.
Holotype: New Zealand. South Auckland. Mangaotaki
Reserve, King Country near Pio Pio. On twigs of
Grtselitnta ltttoralts in deep shade. D.J.Gallo-
WayyeoJune, 1978, \CHR 343163. Isotype in BM.
Thallus lobate-foliose, spreading, in entangled clones,
5-15 (-30) cm diam., loosely attached, corticolous. Lobes
linear-elongate, rather narrow, 3-12 mm wide, expanding
towards apices, + subdichotomously branching, complex, im-
bricate centrally, discrete, + subascendent at apices, mar-
gins entire, slightly thickened below, occasionally with
*The interpretation of P.billardierii (Delise) Rdsdnen
given in Wilkins and James [Lichenologist 11: 274 (1979)]
and in Galloway and James [Lichenologist 12: 293 (1980)]lis
in error and material discussed therein refers to an endemic
taxon which must now be called P.rufovirescens (Church.Bab.)
D.Galloway, comb. nov. Basionym: Sticta richardi var. rufo-
virescens Church.Bab., in J.D. Hook., F1.Nov. Zel.2: 278
(1855).Lectotype: New Zealand. South Island, Akaroa. Hom-
bron, BM ex PC!
206
rounded to elongate white pseudocyphellae, apices truncate
or furcate. Upper surface smooth, glabrous, even or subun-
dulate, rarely very shallowly faveolate, ridges smooth, in-
distinct, without isidia, pseudocyphellae or soredia, con-
spicuously and irregularly white-maculate (x 10 lens) ,dark
Slate-grey to bluish-grey when wet, pale greenish-grey suf-
fused brownish when dry. Medulla white. Phycobiont blue-
green, Nostoc. Lower surface pale brownish-pink, wrinkled-
striate centrally, + evenly tomentose to margins, tomentum
pale, whitish, short, even, lobe apices sometimes glabrous,
whitish, wrinkled-bullate, shining. Pseudocyphellae round
to irregular, common, plane, intense white, 0.1 - 1 - 2 mm
diam., conspicuous.
Apothecia very rare, marginal, subpedicellate, 0.5-2.0 mm
diam., disc glossy, coriaceous, dark chestnut-brown to
black, epruinose, margins pale pinkish-brown, corrugate-
striate to’ verrucose, + inflexed and obscuring disc at tires
becoming + coronate or denticulate to excluded at maturity,
thalline exciple coarsely verrucose-scabrid, minutely to-
mentose, whitish to dark brownish-pink or red-brown. Epi-
thecium 13-22 um thick, yellow-brown, of thickened, conglu-
tinate tips of paraphyses. Hymenium colourless 40-65 um
tall. Paraphyses dense, straight, simple, 1.5 um thick, api-
ces clavate, red brown or yellowish-brown, to 5 um thick.
Asci and ascospores not seen.
Chemistry: 7B-acetoxyhopane-22-0l and hopane-15a-22-diol.
Specimens examined: New Zealand. North Island. South Auck-
land: Kauaeranga River, J.K.Bartlett (Herb Bartlett), Kai-
mai Range, J.K.Bartlett (CHR 343298), Mangaotaki River, D.
J.Galloway (CHR 343277), Hawke's Bay: Road to Lake Tutira,
W.Martin 5484 (CHR 375498), Napier-Taupo Road near Tarawera,
J.K.Bartlett (Herb.Bartlett), Wellington: Erua Swamp, J.K.
Bartlett (CHR 343207), South Island. Nelson: Cobb Ridge, J.
K.Bartlett (CHR 343162), Canterbury: Mt Sinclair Reserve,
Banks Peninsula, D.J.Galloway (CHR 343209), Southland: Fo-
rest Hill, J.Murray (BM).
P.murraytt has a wide distribution in New Zealand
though it is local and rather sparse in presently
known habitats. It is an epiphyte of Dracophyllum
SuDuULaLUM,, GYriseLtinta lLittoralte, Lopnomyrtuenoues
lata, Nothofagus menztestt and Wetnmannta racemo-
sa in areas of high humidity and moderate to dense
shade. It associates with the following lichens:
207
Cocecocarpta erythroxylt, Degelta gayana, Ertoder-
ma neozelandtca, Hypotrachyna stnuosa, Lobarta
Yervgera, 1 .scropiculata, fannarita fulvescens,
Parmelta amphtbola, Letoderma pyenophorum, Physma
chitlense, Polychtdtum contortum, Pseudocyphella-
PLMECUraLa wt CrOCata, sepisTicta,. P.faveolata.,
PMIOCKert, EP atneritcard, © .pstlopny lla... rufovt—
Rescens, P.subvartabtits, Sttcta tatrfrons and
Sbeeta werge lite.
It is closely related chemically and morphologi-
cally to P.rufovtrescens and the two species have
been found attached (joined material comes from
Mangaotaki River (North Island) and Mt Sinclair
Reserve (South Island) only). When growing inde-
pendently it is separable from P.rufovtrescens
in the smoother, undulate, very seldom faveolate
lobes which are expanded at the apices, the
leaden-grey colour when wet because of the pre-
sence of blue-green phycobiont, the consistent
development of a + uniform tomentum on the lower
surface, and the dark brown to black apothecial
discs with corrugate-striate to verrucose margins
and tomentose exciple. In P.rufovtrescens tomen-
tum on the lower surface is very rudimentary and
terestricted to central parts of mature lobes,
IMargins and apices being regularly galbrous, white
and shining. Also the pseudocyphellae of P. rufo-
vtrescens are smaller, more pocklike and scattered
and not conspicuous and large as they are in P.
murraytt. P.murraytt is named for the late Dr
James Murray of Otago University, New Zealand who
first became interested in New Zealand's lichens
through his chemical studies in Pseudocyphellartia.
He was engaged in monographing Stteta and Pseudo-
cyphellarta at the time of his death (1961), and
made the first collections of P.murraytt from
New Zealand in January 1957.
RESULTS
Morphology of phycosymbiodeme development
In the three phycosymbiodemes investigated the
chlorosymbiodeme originates from the cyanosymbio-
deme. In the pair "P.margaretae/P.pubescens", the
208
chlorosymbiodeme (P.pubescens) often arises from
the upper cortex and is formed laminally. However,
marginally developing chlorosymbiodemes and those
Originating from the lower surface of P.margare-
tae may also be observed. By contrast in the pair
"PIMUrrayt] Pvruyovirescene"” “and (°Pinglldan tt) pices
rtacea", laminally developing chlorosymbiodemes
were not seen. Most frequently the chlorosymbio-
demes of these pairs form on the lower cortex and
along the margins. Critical observations of both
cyanosymbiodemes disclose some notable correla-
tions between morphology and the sites of chloro-
symbiodeme emergence:
1. Cyanosymbiodemes with a glabrous upper surface
(P.atlanit ‘and P.rufovirescens) show chilorosym-—
biodeme primordia only on the tomentose lower
surface and along the margins. 2. Cyanosymbiodemes
with a tomentose upper surface (P.margaretae)
have chlorosymbiodeme primordia not only on the
lower surface and margins but also on the upper
surface (figs. 5,6a). The development of chloro-
symbiodemes therefore seems to occur in those
parts of the cyanosymbiodeme thallus where tissues
undergo externally orientated differentiation.
The inoculation of appropriate green algal cells
is evidently restricted to very young hyphae (fig.
4f). The process of encapsulation may be ex-
plained by the accidental close coincidence of
green algal cells and young hyphal cells.Active
outgrowing of hyphal cells triggered by some
stimulus from the algal cells does not appear to
be involved since epiphytic green algal cells may
sometimes cover large areas of both surfaces of
figure 1: Habitat photographs of the pair "P.murrayii/
P.rufovirescens"; a,b:"P.rufovirescens" (chlorosymbio-
deme) bearing young (b) and mature (a) apothecia (ap)
as well as pycnidia (py); c: lower surface of "P..
murrayii" (cyanosymbiodeme)-lobe, with 10 chlorosymbio-
deme primordia formed one beside the other; d: lobe of
"P.,murrayii" bearing a young, stalked lobule of "P.
rufovirescens" which develops marginally; e: lobes of
"P.,murrayii" with young chlorosymbiodemes on the lower
surface; chs: chlorosymbiodeme; cys: cyanosymbiodeme;
scale: imm (e:as in qd).
y yi
Wit
e) #
Ah pe
= + =
“
O.
Hp i
»
either symbiodeme without producing any effect
at all. In all cases studied, chlorosymbiodeme
development begins with the contact of very slight
hyphae surrounding usually only one or a few al-
gal cells (figs. 4f,6c). These tender hyphae or
outermost hyphal cells of the cortex are normal
cortical ‘constituents... The first sstep sinc loro
symbiodeme development, once epiphytic green al-
gal cells are recognized by the mycobiont, is the
immediate surrounding of the algae by hyphae grow-
ing from the cortex of the cyanosymbiodeme (figs.
2£,4c), a process analogous to the early stages
of the development of cephalodia in genera of the
Lobartaceae.
The resulting primordia, regardless of their site
of formation are at first nearly spherical (figs.
3c, 4c) and in them the hayphal envelope forms
from the very beginning a kind of pseudoparenchy-
matous cortex. They later elongate and become
slightly falttened (figs. ;ld,e, 3b,4e) as) the vi-—
sible chlorosymbiodeme. At the apices of these
young lobules an heteromerous arrangement of
tissues is evident. Sometimes also the presumptive
lower cortex may already have developed pseudocy-
phellae (fig. 4e).. During ae periodsof turther
growth, with continuously changing micromorpholo-
gy, a typically organized green-algal-containing
Lobe’ with a distinct upper. ands» lower. cocrex, ne
dulla and algal layer is formed. Whereas thestruc-
ture of the cortical hyphal cells, the thickness
of the cortices, and that of the whole thallus
are Similar in both phycosymbiodemes, at the site
figure 2: Thallus anatomy of the pair "P.murrayii/P.rufo-
virescens"; a: cross section of "P.murrayii" with upper
cortex, algal layer and part of the medulla; b,c: cross
section of "P.rufovirescens" with upper cortex, algal
layer and part of the medulla, b: young lobe in close
Vicinity co “P.murrayii", c: mature lobe;i¢,e:) cross
section of "P.rufovirescens", showing internal cephalo-
dia (ic) developing from the lower thallus surface (d)
and exeptionally from the upper thallus surface (e);
f: cross section of "P.murrayii" with one chlorosymbio-
deme primordium (chsp) on the lower cortex (lc); scale
in c (representative also of a,b,e,f) and in dad: 20 um.
Pa Ml
Pvp
Hi Hib iii i) iif
i}
i i ‘ ,
da
i ha. a
Wl ise)
Za?
of attachment these anatomical structures may
successively change with continuing growth of the
chlorosymbiodeme and its increasing distance from
the cyanosymbiodeme. This drifting apart of deve-
lopmental morphology is best seen in the pair
"P.murraytt/P.rufovirescens".
At the base of the thallus of P.rufovtrescens
which is very close to the thallus of VP. murray,
the cells of the cortices are + isodiametric with
large lumina and rather thin cell walls, resem-
bling: those of Pimurraytt (fig.2b). sine green var.
gal cells are arranged in a wide-ranging layer
with numerous aplanosproangia and free aplanospores
accounting for the small size of the algal cells
in this region. The green algal layer reaches deep |
into the medulla and sometimes nearly touches the
lower cortex. If the chlorosymbiodeme is attached
to the cyanosymbiodeme by a stalk-like transition
zone (fig 1d), then this area is devoid of algae
and the formation of the upper cortex of Jeheschlo-—
rosymbiodeme is seen as a sudden constriction in
the cortex of the stalk. With increasing distance
from the chlorosymbiodeme's growing tip, the al-
gal layer becomes confined to a thin zone beneath
the upper cortex (fig. 2c), the algal cells being
closely packed and only rarely developing aplano-
sporangia. The upper cortical cells are no longer
exclusively isodiametric (fig. 2c) .The outer layer
especially consits of periclinally orientatedthy—
phal elements with small cell lumina and thickened
cell walls. The lower cortex develops a sparse
tomentum. In this zone internal cephalodia may de-
velop (fig. 2d). Exceptionally, cephalodia origi-
nating from the glabrous upper surface are formed
(fig. 2c). In contrast to the cyanosymbiodeme,the
chlorosymbiodeme bears both apothecia and pycni-
figure 3: Habitat and SEM-photographs of the pair "P.
allanii/P.coriacea"; a: greater part of "P.allanii"
(cyanosymbiodeme)-thallus with lobules of "P.coriacea"
(chlorosymbiodeme); b: young lobule of "P.coriacea" de-
veloping marginally from "P.allanii"; c: SEM-photograph
of a chlorosymbiodeme primordium on the lower cortex of
"P,allanii" surrounded by hyphae of the tomentum.
213
ht
ji
214
dia» ia gz tay, De.
An inhibitory effect of the developing chlorosym-
biodeme in suppressing the formation of further
primordia in its immediate vicinity was not demon-
strated. Sometimes one may find up to ten chloro-
symbiodeme primordia growing close together (fig.
1¢),+a Lact, rendering unlikely the, action ofeany
soluble morphogen. The lower cortex of the chloro-
symbiodeme does not undergo any remarkable diffe-
rentiation and is therefore not different from
that of the cyanosymbiodeme. The same is true for
pseudocyphellae which appear, in the cyanosymbio-
deme, to be places of heightened secondary meta-
bolic activity; they are nearly always filled
with large amounts of lichen substances.
In order to show the influence of either algae on
the formation of particular structures in the phy-
cosymbiodemes of one pair, the development of
pseudocyphellae was studied. These are characte-
ristic of species of Pseudocyphellarta. The re-
sults accord with those of Renner (1980) for P.
freyctnetit and P.thouarsit. The first discernable
sign in the process of formation of pseudocyphel-
lae is the appearance of typically differentiated
medullary hyphae in a circumscribed region of the
medulla directly above the lower cortex. These
figure 4: Thallus anatomy of the pair "P.allanii/P.coria-
cea": a,c,d,e,£: cross sections of "P.allanii”™; a: ana—
tomy of upper cortex and algal layer; c: chlorosymbio-
deme primordia (chsp) growing from the lower cortex (lc)
of "P.allanii"; d: chlorosymbiodeme growing with a stalk
(st) on the lower surface of the cyanosymbiodeme: note
that the stalk is devoid of algal cells; the algal-con-
taining tip is slightly separated from the stalk by a
constriction (arrows); e: very young lobe of "P.coria-
cea" in which a prospective upper cortex (puc) and pro-
spective lower cortex (plc) may be discerned, the latter
having already developed pseudocyphellae (ps); f£: very
early stage of chlorosymbiodeme formation, showing very
tender hyphae of the tomentum (shy) surrounding a few
algal cells; b: cross section of "P.coriacea" with upper
cortex and algal layer; scale in c (representative also
Of «a,b, 1e (aS. Aud) ere COLO.
215
216
hyphal initials consist of branches with short,
swollen cells with enlarged lumina(fig.7a). Some
of these cells make contact with the lower cortex
and) forcibly, cupturevit. While growing, threudch
the cortex the hyphal initials further subdivide
into many short-articulate cells (fig.7b). The
lower cortex first invaginates and later disinte-
Grates, probably, through. gradual lysism(tige/c)e
Once the cortex is breached, the medullary hyphae
are in free contact with the substrate. Further
enlargement of this opening in the cortex? is \ini=
tiated along the border of the pseudocyphellae
(fig.7c). In some cases the lower cortex bordering
the pseudocyphellae become + crateriform. In this
general scheme of development no differences be-
tween species of Pseudocyphellarta with green al-
gae and those with blue-green algae can be shown,
nor are there differences between the two phyco-
symbiodemes. Nevertheless the final morphological
appearance of these structures, e.g. the frequen-
cy, diameter and shape of pseudocyphellae may
vary to some extent as is true for cyphellae in
Stteta (Renner 1980).
The course of chlorosymbiodeme development out-
lined torn the pair "P.murrayi7/P.rupovireecens.
LS similar. tor that! of ‘the otherstworparrs Ofspoy—
cosymbiodemes studied (figs. 3,4,5,6). Although
Similarities in the development of phycosymbio-
demes discussed by James and Henssen (1976) in
DEvera, andwine rel crgera by, Brodoyandwkichardson
(1979) are obvious, one striking feature distin-
guishes the pairs of phycosymbiodemes in Pseudo-
cyphellarta from these last, viz., the remarkable
resemblance of both phycosymbiodemes forming a
pair. When dry they are scarcely distinguishable
from each other and when wet their morphologies
are closely similar but thallus colour bstystri-
kingly different. Clear evidence of different
figure 5: Habitat photographs of the pair "P.margaretae/
P.pubescens"; b: greater part of "P.maragaretae" (cyano-
symbiodeme) bearing numerous leaflets of "P.pubescens"
(chlorosymbiodeme); a: enlarged part of b, giving evi-
dence of the hairy upper cortex of both, the cyanosym-
biodeme (cys) and the chlorosymbiodeme (chs).
Ze.
ZS
thalli distinguished only by the presence of two
kinds of algae is only obtained after cutting the
lichen sample.
Chemistry
Fragments of phycosymbiodeme pairs were mechani-
cally isolated under microscopic control and then
extracted in cold acetone and chromatographed.
Results of ascending TLC in solvent system C are
given (tig.9)"\ (For “Pomardaretae/P,puveseencs,
two-dimensional TLC was necessary to demonstrate
the occurence of methyl lecanorate and methyl
evernate and of norstictic acid (fig.10). Neutral
compounds belonging to the main substances were
further characterised by their mass spectra. De-
tailed data are presented elsewhere (Renner 1980).
In two pairs of phycosymbiodemes
Viz., “P.murraytt/P. rufovtrescens", and "Pvavlani 7,
P.cortacea" results give strong evidence of two
triterpenoids, 7B-acetoxyhopane-22-ol and hopane-
15a-22-diol (Wilkins and James 1979, Renner 1980)
being present in both joined thalli as well as in
either phycosymbiodeme (fig.9). Mass spectral da-
ta of 7B-acetoxyhopane-22-ol:
M : m/e 486 (not registrated), m/e 468, m/e 453, m/e 428
(not registrated), m/e 393, m/e 249, m/e 206, m/e 205,
m/e 207, m/e 189 and m/e 149.
Mass spectral data of hopane-15a-22-diol:
M : mfe 444, m/e 426, m/e 411, m/e 386, m/e 223, m/e 205,
m/e 191 and m/e 165.
The chemistry of the pair "P.margaretae/P.pubes-
eens" is more complex, containing substances from
figure 6: Thallus anatomy of the pair "P.margaretae/P.
pubescens"; a,c,d: cross sections of "P.margaretae"; a:
upper cortex and algal layer; c: chlorosymbiodeme prim-
ordium on the upper cortex, containing two green algal
cells(a) being intimately surrounded by hyphae of the
tomentum; d: lower thallus cortex with part of the me-
Gulla; the structure of the lower cortex is the same as
in "P.pubescens"; b: cross section of "P.pubescens"-lobe
in proximity of "P.margaretae" with upper cortex and ad-
joining algal layer; scale in d (representative also of
asi ge) soo20) Lin.
219
SS
=
=
220
all three main biosynthetic pathways involved in
secondary metabolism in lichens. Among the neu-
tral constituents, hopane-15a-22-diol and hopane-
60a-—/B=22—triol were clearly identifticdw(fig. 97.
Mass Spectral; data Of nopane-6a—-7b-27-curo0l::
M<:7m/e 460, m/e 442, m/e 427, m/e 402, m/e 223, -m/e.zC07,
m/e 189 and m/e 149.
These two triterpenoids of the hopane-series are
present in both phycosymbiodemes(fig. 9,10). There
is possibly also a trace of 7B-acetoxyhopane-22-ol
present in both and one further neutral compound
(triterpenoid, or sterol 7?) \of.fa nignereR ero
ducts of the shikimic acid pathway, icalyvcin pula
vinic dilactone and pulvinic acid are present in
both phycosymbiodemes. Depsidones are norstictic
acid, wSstictic acid, cONnSticticracidwancdmer wy com
stictic acid, with all compounds present min born
thalli. Two tridepsides, tenuiorin and methyl gy-
rophorate (only in trace amounts) are present in
both phycosymbiodemes. The identification of all
other depsides remains speculative - they are
present only in trace amounts and most likely re—
present the additional spots, "Often occuringsra.
gether with tenuiorin and methyl gyrophorate, viz.,
methyl evernate, methyl lecanorate and possibly
two others. Of the four last-mentioned constitu-
ents at least one is not present in P.pubescens.
Among volatile compounds of the methylamine se-
ries, only trimethylamine was detected by gas
chromatography. Aliquots of aqueous extracts of
both phycosymbiodemes yielded similar amounts of
figure 7: Schematic outline of pseudocyphellae-development
in P.thouarsii (taken from Renner 1980);a: very early
stage in the development showing strong coloured hyphae
in contact with the lower thallus cortex (lc) and indu-
cing cells of the cortex into active. growth; b: stage to
be characterized as an invasion of the lower cortex by
articulated hyphal elements; the cortex is ruptured by
the pressure of growing hyphae and lysis (ly); c: young
pseudocyphella, with hyphae directly exposed to the
substratum.
=
Cre 3
Ge
y, Ne pF
|
am as
Qe
OC
0
Os
USS
oe.
oO
Ooo
Zon
222
this compound.
DISCUSSION
The similarity shown by pairs of phycosymbiodemes
Of Pseudocyphellarta studied in the developmental
processes leading to the formation of chlorosym-
biodemes is very striking. In all cases studied
chlorosymbiodemes are secondarily produced by the
mycobiont of the cyanosymbiodeme. Leaving aside
differences in the organisation of other cyano-
symbiodemes as well as different sites of origin,
the sequence of morphogenetic steps in chlorosym-
biodeme development is much the same in Pseudocy-
phelttaria, Stteta (e.g. "S.dufourtt/S.canartensic »
and Pelttgera (unnamed (P.avenosa Gyelnik) [P.W.
James, pers.comm.]blue-green morphodeme with Co-
eecomyxa-containing lobes). Whenever a chlorosym-
biodeme is formed it is of secondary origin and
is characterised by an heteromerous arrangement
of tissues. A schematic representation of this
process is given inviig. 11, with the chlorosym-
biodeme showing heteromerous arrangement of tis-
sues.
Changes in morphology and anatomy in both thalli
of joined species resulting from the morphogene-
tic influences of two different algae vary in
their expression. Differences in both secondary
metabolites, and developmental morphology of
structures such as pseudocyphellae and cyphellae
(Renner 1980) in most cases are negligible. That
both phycosymbiodemes may produce fruiting bodies
(as in an unnamed pair of phycosymbiodemes from
Stteta - Renner 1980) contrasts with assertions
figure 8: Semi-schematic outline of internal cephalodium-
development in Sticta latifrons (taken from Renner 1980);
The course of development starts with the capture of
Nostoc-algae (Al) by fungal hyphae of the lower cortex
(lc), stage a, followed by successive surrounding of al-
gal cells by hyphae, stage b and c, and the invagination
of algal cells into the thallus medulla by hyphal pres-
sure (PHY), stage d, to form an internal cephalodium
which then enlarges within the medulla, stage e,f, is
typical for foliose green-algal lichens in the Pelti-
gerineae.
224
that sexual reproduction might be influenced by
the algae present (James and Henssen 1976).
In the Peltigerineae many chlo-
rosymbiodemes known to occur as independent green-
algal lichens (all chlorosymbiodemes studied her@
contain internal cephalodia as sometimes do the
chlorosymbiodemes themselves. A comparison be-
tween cephalodium development and chlorosymbio-
deme formation demonstrates certain parallels.
Formation of cephalodia may be regarded as the
reverse process to that of chlorosymbiodeme for-
mation, e.g. the secondary lichenisation of blue-
green algae (Wostoc).
The process - illustrated here for Stteta latt-
frons; according to Renner W(1980) j= "startsiwien
capture and encapsulation of epiphytic blue-green
algae (fig. 8a) by hyphae of the mycobiont. Very
early stages in this process show a small, nodule-
like protuberance on the lower surface of the
green phycobiont-containing lichen (or chlorosym-
biodeme) resembling in structure primordia seen
in the formation of chlorosymbiodemes (fig. 8b,c).
Unlike the development of chlorosymbiodemes, fur-
ther differentiation of cephalodia is character-
ised by an invagination of Nostoec-algae into the
thallus medulla (figs. 8d,e,f) or, in the case of
external cephalodia, by a moderate enlargement of
the protuberance sto form a+ sacculate; cori at
hump with cells of WNostoe irregularly distributed
throughout the mycobiont tissue (See Henssen and
Jahns 1973). In no case is a new heteromerous
thallus formed.
In cephalodia the lichenisation of a secondary al-
ga results inva marked change in) metabolitespar=
tern. The same situation obtains in Pseudocyphel-
larta and in Lobarta (Lobarta ampltsstma, Renner
1980, 1982a,b)° and will probably also; bevturucrecr
other members of the Peltigerineae. The change
noted is an absence of secondary metabolites from
the core of the cephalodia. The absence of these
substances may be explained by the diversion of
products of primary metabolism (nitrogen fixation)
of the alga to the mycobiont where they are absor-
bed, and probably subsequently fixed and intercon-
verted (Feige 1976, Renner 1980). These latter
220
figure 9: Thin layer chromatogram (solvent system C acc.
Popculperson, 1972) of °crude Pichen extractsrof all pairs
of phycosymbiodemes investigated here; position 1: "P.
margaretae"; position 2: "P.pubescens"; Position 3:"P.
murrayii"; position 4:"P.rufovirescens"; position 5:
ePLablaniils: positionke: P,corlacea’ ; position 7: stic=
tic-acid-complex standard; position 8: usnic-acid stan-
dard; SF: solvent. front; SL: start line; PDL: pulvinic
dilactone; C: calycin; P: pulvinic acid; T: tenuiorin;
MG: methyl gyrophorate; aH:7B-acetoxyhopane-22-01; NS:
norstictic acid; Hd: hopane-15a-22-diol; Ht: hopane-60-
7B-22-triol; UWS: (Unknown S-2 with Stictic acid sens.
Culberson 1972) identical with cryptostictic acid; CS:
COonsticric acid: Ss stictic acid; Us: usnic acid:
226
IT Dim. (102A) —>
Il Dim. (HEF) —
figure 10: Two-dimensional thin layer chromatogram (sol-
vent system A: 1.dimension, B: 2. dimension, A,B acc.to
Culberson (1972)) of the acetone extracts of "P.marga-
retae"; ML: methyl lecanorate; ME: methyl evernate; all
other abreviations as in fig.9
processes remain to be conclusively proved. Lea-
ving aside the vexed problems involved in demon-
strating the presence of lichen substances in
different parts of the thallus, the question re-
mains to what extent secondary metabolites may
correlate with the alga present. Any discussion of
this question must necessarily at present be spe-
culative. As long as we have no reliable evidence
for any correlative links between the phycobiont's
Ley
metabolic activities and the production of lichen
substances within the mycobiont we are far from
understanding the possible directing role of phy-
cobionts in the expression of phytochemistry in
lichen symbioses. The absence or the additional
presence of some secondary products in either
phycosymbiodeme may well be known but cannot be
determined by considering the algae alone. The
converse case, viz., taking the identity of se-
condary products in both phycosymbiodemes as
DrOOoLt for the continuity, of the mycobiont in’ yoin—
ed thalli (Jordan 1972) is also inadmissable. The
only metabolites known at present to be clearly
confined to the mycobiont are the methylamines
( Bernard et al. 1974, 1980), however these pro-
ducts are of restricted occurence. James and
Henssen (1976) record that Sttcta dufourit has a
distinct fishy smell (methylamines) whereas 5S.
canartensts does not have any smell at all. Con-
trasting with this assertion one of us was able
to prove by gas chromatography (Renner 1980) that
free-living specimens of either species mentioned
above do contain trimethylamine. The proportion
in the two species could not be determined.
Evidence for divergent metabolic activities in
the tissues of lichenised mycobionts comes from
an wn=—sttu localisation of lichen acids using a
cytophotometri¢ technique (Renner 1982a,b). It is
shown in this study that in contrast to high con-
centrations of depsides and depsidones in the
medulla, no substances of this class were detec-
table within the fungus tissue of cephalodia in
green-algal species of Pseudocyphellarta, Lobarzta
and Nephroma. Further, the detection in some
species of Pseudocyphellarta of sustantial con-
centrations Of secondary products in particular
thalline structures (e.g. pseudocyphellae and
soralia: in "P.margaretae/P:pubescens" the yellow
pseudocyphellae contain pulvinic acid and its de-
rivatives) or in the medulla, supports the assump-
tion of a heterogenous metabolic activity of the
lichenized mycobiont. These facts cannot at pre-
sent be satisfactorily explained.
The reflections outlined here have some important
implications for interpretations based on secon-
228
dary metabolites detected in both phycosymbio-
demes or in structures of the green-algal thallus
containing blue-green algae, e.g. cephalodia.
Whereas in the inner mycobiont tissues of some
external cephalodia such as in Nephroma kuehne-
mantt , no metabolites are detectable, the cor-
tex of the cephalodia may give a positive result
for at least some secondary metabolites charac-
teristic of Wiokuehnemant~7, (Renner 1950, 19382b)e
Extraction of excised cephalodia of this lichen
and analysis of the extract while showing the
presence of secondary metabolites, does not prove
that the detected compounds are synthesised in
the cephalodia under the influence of the secon-
dary algae (Wostoc).
The difficulties inherent in correlating secon-
dary metabolites with any particular algae in the
symbiotic association will only be resolved by
carefull labelling experiments with C-labelled
precursor substances and such investigations are
Curently win progress.
figure 11: Schematic outline of phycosymbiodeme forma-
tion in Pseudocyphellaria; a: blue-green algal lichen;
if a tomentum (t) is present on the upper cortex (uc)
and on the lower cortex (lc) the capture of green algal
cells may occur’ on both .cortices; chspsichlorosymbics
deme primordium; ap: apothecia; all: algal layer; m:
medulla; psc: pseudocyphellae; b: resulting pair of phy-
cosymbiodemes, with lobes of the chlorosymbiodeme (chs)
on either surface of the cyanosymbiodeme (cys) at sites
of formation of chlorosymbiodeme primordia; above: fer-
tile chlorosymbiodeme with tomentum on both cortices
and the development of internal cephalodia (dic); the
fully developed chlorosymbiodeme may represent a com-
bination of the two demonstrated alternatives.
Cia)
a : Eo ! He ees ee
230
ACKNOWLEDGEMENTS
We are grateful to J.K. Bartlett (Pakuranga,
Auckland) for the provision of specimens, to Dr
Elizabeth “tbdgar. i) Botany Division yeD~S-dens,
Private «Bag, Christchurch) fon -thesbatineaiad=
noses, and to P.W.James (BM) for critically rea-
ding the manuscript and making many constructive
comments upon it.
Rech Se. tent, Ciews
BERNARD,T.,M.GOAS & P.L.MAILLET: Sur le systéme va-
cuolaire a granules argentafines des lichens a trimé-
thylamine de la famille Stictacées.J. Microscopie
LOM NSO. 3 ound. owns)
BERNARD,T.,M. JOUCLA, G.GOAS et J.HAMELIN: Carac-—
térisation da la sticticine chez le lichen Lobarta lae-
tevtrens. Phytochem. 19, 1967-1969 (1980)
BRODO,I.M. & D.H.S.RICHARDSON: Chimeroid association
in the genus Pelttgera. Lichenologist 10, 157-170 (1979)
CULBERSON,C.F.: Improved conditions and new data for
the identifictaion of lichen products by a standardized
TLC, method. J.Chromatoga/2, 1N3—i25 (1972)
DUGHI,R.: Une céphalodie libre lichénogéne:Le Dendrisco-
cauton bolanteum Nyl. Bull.Soc.bot.Fr.84, 430-437
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DUGHI,R.: Sur les relations, la position sytématique et
l'extension du genre Densriscocaulon. Annls. Fac.
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FEIGE,B.: Untersuchungen zur Physiologie der Cephalodien
der Flechte P¢ ttgera aphthosa(L.)Willd. II Das photo-
synthetische C-Markierungsmuster und der Kohlenhydrat-
transfer zwischen Phycobiont und Mycobiont. Z.Pflan-
Z enphysiol. 80,386-394 (1976)
HENSSEN, A. und H.M.JAHNS: Lichenes, Eine Einfthrung
in die Flechtenkunde. Thieme,Stuttgart 1973 ["1974"]
JAMES,P.W. & A.HENSSEN: The morphological and taxono-
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Lichenology: Progress and Problems (D.H.BROWN,
D.-b HAWKSWORTH & ROH -BAILEY -eds'.)eAcademi cy yres:
1 OG
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Lee Leta po haa, CI97 5)
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JORDAN,W.P.: Erumpent cephalodia, an apparent case of
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MYCOTAXON
VOL AVI INO depp oe =258 October-December 1982
A Note on Sporotrtchum gougerottt Matruchot 1910
Michael R. McGinnis and Libero Ajello
Department of Bacteriology and Immunology,
University of North Carolina at
Chapel Hill, Chapel Hill, North Carolina 27514 and
Division of Mycotic Diseases, Center for Infectious Diseases,
Centers for Disease Control, U.S. Public Health
Service, Department of Health and Human Services,
Atlanta, Georgia 30333, U.S.A.
SUMMARY
The taxon Sporotrtechum gougerottt Matruchot 1910 is
rejected as a nomen dubtum because the name is of uncer-
tain application. Owing to the absence of type material,
an inadequate description, and the lack of illustrations,
there has been no consistent taxonomic concept for this
fungus. We reject as invalid the recently proposed new
combinations of Exophtala gougerotit Ravisse et Rodriguez
Vindas 1981, FE. gougerottt South, Brass et Stevens 1981,
and #. gougerotit Nishimura et Miyaji 1982.
INTRODUCTION AND DISCUSSION
In 1907, Beurmann and Gougerot (1) described an
atypical case of sporotrichosis (case XI) in a patient
suffering from tuberculosis. The infection manifested
itself as a painless, deep, hard nodule on the right thigh.
Prolonged direct examination of pus aspirated from the
nodule revealed a single, oblong fungal form that Beurmann
and Gougerot considered to be compatible with that produced
by a species of Sporotrtchum Link. The fungus that was
isolated from the aspirated pus was described as being
black, slow growing, and producing bouquets of 6 to 12
spores with a few spores being borne singly only in the
Zoo
center of the colony. On the basis of these findings, the
authors equated their isolate with Sp. beurmanntt (as
beurmannt) Matruchot et Ramond 1905, which is now con-
sidered to be a later synonym of Sporothritx schenektt
Hektoen et Perkins 1900. Unfortunately, Beurmann and
Gougerot paid little attention to the alleged tissue form
of the fungus, which cast doubt on the very existence of
a mycotic infection. On the basis of the data furnished
in their case report, one could conclude that the recovered
fungus was merely a contaminant.
Beurmann and Gougerot gave their isolate to Matruchot,
who in 1910 (13) described it as Sp. gougerotit (as
gougerott). It is uncertain whether or not Matruchot
intended the name gougerotit to be used as a variety or
species name. On page 545 of his paper (13), he wrote
"Je le considere comme une variete stable du S. Beurmannt
et le dénomme S. Gougeroti". Later on that same page,
Matruchot stated that among the small group of fungi that
caused sporotrichosis, he recognized Sp. gougerotit as one
of the "...trois types assez differents pour constituer
trois especes distinctes ...'"' In one instance he proposed
Sp. gougerottt as a variety of Sp. beurmannitt, and in a
second instance a distinct species. Since Matruchot did
not refer to Beurmann and Gougerot's description of the
isolate presented to him (1), the name Sp. gougerottt must
rest solely upon the description that Matruchot provided
in his study. Matruchot considered the aerial fructifi-
cations produced by Sp. gougerottt to be intermediate to
those of S. schencktt and Sp. beurmannit. In addition,
Matruchot described a "budding" form as being character-
istic of Sp. gougerotit. Judging from his description,
the "budding" form may have consisted of, in part, toruloid
hyphae. The description provided by Matruchot is so vague
that it is impossible to determine the diagnostic char-
acteristics of his fungus, which has resulted in substan-
tial taxonomic confusion. Unfortunately, a specimen was
not prepared, illustrations were not provided, and the
culture that Matruchot had used for the description of
Sp. gougerottt was not saved. As a result, Matruchot's
taxon has been classified in the genera Rhtnocladtum
Saccardo et Marchal (22), Torula Persoon (12), Demattum
Persoon (7), Oospora Wallroth (10), Cladosporium Link (5),
Phtalophora Medlar (2), and Rhtnocladtella Nannfeldt (19)
by various investigators.
234
Castellani and Chalmers (6) considered Sp. gougerottt
to differ "....from the typical Sp. beurmannii only in
small details, the principal ones of which are the black
pigmentation of the colonies from the very beginning, and
the extremely abundant sporulation."" This disposition can
be supported by examining the exquisite drawings prepared
by Nannizzi (16) for figure 194 in his text book. These
show the characteristic sympodial conidiogenous cells and
arrangement of conidia that are typical of the genus
Sporothrix Hektoen et Perkins. In contrast, Gritz (8) and
Grigoraki (7) characterized Sp. gougerottt as producing
what appears to be toruloid hyphae with and without clusters
of round cells associated with the hyphae. Some of their
illustrations are suggestive of Montltella Stolk et Dakin.
Janke's (10) concept parallels that of Grutz, with the
additional morphological form of budding yeast cells being
included (see his figures 6 and 11). A third interpre-
tation of Sp. gougerotit was presented by Kesten et al.
(11), Young and Ulrich ’ (23) ,. and Borelli. (3), who depicted
the conidiogenous cells as being cylindrical and bearing
clusters of conidia at their apices as seen in the genus
Exophtala Carmichael.
In 1953, Young and Ulrich (23) isolated a dematia-
ceous hyphomycete from a human case of phaeohyphomycosis.
They identified it as Sp. gougerotit and sent it to
Gougerot, who confirmed their identification. Borelli
obtained a subculture of Young and Ulrich's (NCMH 575)
isolate and in 1955 designated it as the neotype for Sp.
gougerotit (3). He then transferred the taxon to the genus
Phtalophora (2). Borelli considered FH. jeanselmet (Langeron)
McGinnis et Padhye 1977 and Wangtella dermattttdts (Kano)
McGinnis 1977 to be later synonyms of P. gougerotit.
Because a living culture can not serve as a neotype
(Article 9.5, ICBN) (21) and a specimen was not provided
(4), Borelli's proposed neotype is invalid. In addition,
even if his proposed neotype had been prepared properly
as stipulated under the rules of nomenclature, his neotype
could not be accepted because it can not be shown that the
neotype is compatible with the protologue of Sp. gougerotit
(CATti Cre 13215) LCEN)..
The problem of Gougerot's confirmation (23) of Young
and Ulrich's isolate as Sp. gougerotit needs to be taken
under consideration. It is important to remember that the
Pikes
protologue fixes the characteristics of the taxon. NCMH
575 could have been typical of the culture that Gougerot
gave to Matruchot, but it may not have been what Matruchot
actually described. In addition, Matruchot (13) made no
reference in his paper to the description of the fungus
that Beurmann and Gougerot provided in their paper (1).
If Gougerot and Matruchot had saved their original cultures
or specimens, this doubt would not have arisen. Even
though Gougerot considered NCMH 575 to be Sp. gougerotit,
a fungus that he had isolated 46 years earlier, the des-
cription provided by Matruchot must be followed. Based
upon the descriptions provided by Beurmann and Gougerot
and by Matruchot, it is now impossible to determine if
these authors were working with the same fungus.
In 1968, Schol-Schwarz (19) proposed the new combi-
nation Rhinocladtella mansontt (Castellani) Schol-Schwarz.
She treated Sp. gougerotit as a later synonym of this
taxon. Recently, de Hoog (9) considered Sp. gougerotit
to be a later synonym of £. mansontt (Castellani) de Hoog
1977. McGinnis (14) concluded that Sp. gougerotit sensu
Borelli represents misidentified isolates of EF. jeanselmet
and that Sp. gougerottt sensu Matruchot was a later
synonym of S. sechencktt.
In three recent independent studies, the new combinat-—
tion #. gougerottt has been invalidly proposed. Ravisse and
Rodriguez Vindas (18) in 1981 cited "Exophtala gougerottt
(Phtalophora gougerottt), McGinnis et Padhye, 1977" as an
etiologic agent of phaeohyphomycosis. Because McGinnis
and Padhye (15) did not propose this combination, the name
E. gougerottit must be attributed to Ravisse and Rodriguez
Vindas. This name is invalid and must be rejected because
they did not publish it in accordance with Article 33.2
of the ICBN. South, Brass, and Stevens (20) also invalidly
proposed the name £. gougerotitt and incorrectly implied
that Zaias (24) had used this name. Nishimura and Miyaji
(17) in their studies on W. dermatittdts used the binomial
E. gougerotit. As in the case of Ravisse and Rodriguez
Vindas, the latter two proposals must be rejected for the
same reason. The invalid name F. gougerottt could have
been avoided, if these three groups of investigators had
followed the provisions of the International Code of
Botanical Nomenclature.
250
In summary, we believe that the name Sp. gougerottit
should be rejected because the identity of Matruchot's
fungus can not be determined with certainty. The name
Sp. gougerotitt is considered to be a nomen dubium
because its application is uncertain. This taxonomic
problem exists because the original cultures and specimens
were discarded, illustrations were not provided, and the
description of the taxon was inadequate.
ACKNOWLEDGEMENTS
The authors wish to thank Dr. Donald Rogers for
reviewing our manuscript.
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258
Code of Botanical Nomenclature Adopted by the
Twelfth International Botanical Congress,
Leningrad, July 1975. Bohn, Scheltema and
Holkema, Utrecht.
eA eec sity Wee | aie Rhee, Précis de Parasitologie Humaine.
Ind ea.i0.y Doin, Paris.
23° Young, JaM.. and Ee Ulrich. 1953. Sporotrichosis
produced by Sporotrichum gougerott. Report of
a case and review of the literature. Arch.
Dermatol. Syphil (Chicago) 67:44-52.
24, Zaias, N. 1978. Chromomycosis. J. Cut. Pathol:
5:155-164.
MYCOTAXON
VOUee we Noy iy pp, 32592242 October-December 1982
et
PARAFFIN EMBEDDING AND SEMITHIN SECTIONING
OF BASIDIOCARP TISSUES
KELLY A. GRATTO and DARRYL W. GRUND
Department of Biology
Acadia Universtty
Wolfville, Nova Scotta, Canada
BOP 1lxo
ABSTRACT
Microscopic details of basidiocarps are important in species
determination among agarics. A method of embedding in paraffin
was devised to demonstrate sporocarp microscopic morphology.
Semithin (3 um) microtome sections were obtained that clearly
revealed the cuticle, context, and tramal region; the hymenium
remained indistinct.
INTRODUCTION
Microscopic characters are important to taxonomic studies, especially
at, or below, the species level (Singer, 1962; Hesler & Smith, 1979).
Agaricologists typically have used hand sectioning to study the micro-
scopic morphology of basidiocarps. Paraffin sectioning should provide
more consistent results, but no satisfactory technique has been
reported. The technique described here produces material which can
be cut to 3 um thickness. The uniform quality and thinness of the
sections make them well suited for photography. As an additional
advantage, the permanently prepared slides can be kept for future
reference.
MATERIALS AND METHODS
Pieces of the pileus and stipe of fresh basidiocarps were placed in
aJ0:25:5 (v/v) water, ethanol, formalin fixative (Corner, 1967) for
a minimum of 24 h. Portions approximately 4x4 mm were cut from these
pieces and dehydrated in an upgraded series of tertiary butal alcohol
(TBA) solutions (Johansen, 1940): TBA 1 (2 h), TBA 2 (2 h), TBA 3
(2 h), TBA 4 (2 h), TBA 5 (overnight), and finally absolute TBA (3
changes; 4 h, 4 h, and overnight). Following dehydration, the
tissues were infiltrated by three 20-minute changes of molten
paraffin (56°-57°C), and embedded in fresh paraffin. Tissue blocks
240
were sectioned at 5 and 3 um using a retary microtome and sections
mounted on slides lightly coated with Mayer Albumin Fixative (Humason,
1979). Sections were floated on a thin film of distilled water
applied to the edge of the ribbon. The slides were transferred to a
39°C warming tray for 6-12 h, deparaffinized in two changes of xylene
(5 and 3 min respectively), and rehydrated by means of a downgraded
series of ethanol: 100% and 95% (5 min each); 85%, 70% and 60% (3
min each). The slides were further rehydrated in 3% (w/v) potassium
hydroxide (KOH) (5 min), stained in a 1:1 (v/v) aqueous solution of
1% phloxine and 1% Congo Red (30 min), rinsed in distilled water, and
again placed in 3% KOH (15 min). Coverslips were mounted using
Kaiser Glycerine Jelly (Humason, 1979).
RESULTS
Sections were cut at 5 um thickness to demonstrate the cuticular
layers and at 3 um to show the trama and context. Satisfactory
sections of less than 3 um in thickness could not be obtained as
tissues tended to compress and wrinkle during sectioning.
The 3 um thick sections clearly demonstrated the pileal and
stipe cuticles, context, and lamellar trama (Figs vl. 3 See yeep
morphology of the hymenial elements was indistinct. Despite the lack
of structural detail in the hymenium, it was possible to determine
the relative density and positioning of the cystidia (Fig. 4).
DISCUSSION
While hand sectioning is a useful method, paraffin embedded
tissues yield more consistent results as they can be cut at a uniform
thinness.
Berta (1976) described an embedding method using butylmetha-
crylate and styrene with benzoyl peroxide from which 2 um thick
sections of agaric tissues were obtained. With respect to morphology,
the 3 um thick sections cut from paraffin embedded tissues are com
parable to Berta's (op. cit.) results. Plastic embedding techniques
involve potentially carcinogenic, costly, and non-reusable materials
in contrast to the less toxic, less expensive, and reusable chemicals
involved in the paraffin method.
ACKNOWLEDGEMENTS
This work was financed in part by an N.S.E.R.C. Postgraduate
Scholarship.
Figures 1-4. 1. Stipe cuticle of Lactarius vellereus (section
5 um thick). 2. Pileal cuticle of L. volems (section 5 ym thick).
3. Context tissues of L. deceptivus (section 3 um thick). 4. Trama
and hymenium of LZ. volemus (section 3 wm thick).
241
242
REFERENCES
Berta, G. 1976. A contribution to the procedures for semithin
sections for critical tissues. Allionia 21: 59-65.
Corner, E.J.H. 1967. A monograph of Clavarta and allied genera.
Oxford Univ. Press, London. 740 pp.
Hesler, L.R. and A.H. Smith. 1979. North Amer. species of
Lactarius. Univ. of Mich. Press, Ann Arbor, Mich. 841 pp.
Humason, G.L. 19/79. Animal tissue techniques, fourth ed.
W.H. Freeman and Co., San Francisco. 661 pp.
Johansen, D.A. 1940. Plant microtechnique. McGraw-Hill Book
Con, 2ines., News.orkie /525.pp.
Singer, R. 1962. The Agaricales in modern taxonomy. J. Cramer,
Weinheim. 915 pp.
MYCOTAXON
Vor AVED NO. Li spp 2455248 October-December 1982
BASIDIOSPORE VARIATION IN LOCAL POPULATIONS
OF SOME APHYLLOPHORALES
ERAST PARMASTO & ILMI PARMASTO
imstttute: of 200l0gy and Botany
202400 Tartu, Estonian SSH, USSR
The variability of mean spore length, width and spore form
index @ (mean length divided by mean width) of eleven local
populations of five Phelltnus spp. and one local population of
Polyporus rhtzophtlus was studied. The coefficient. of variation,
V, for spore length and width usually lay between 2-10, and for @
between 1-8. These values are within the range found for similar
parameters in other organisms.
The variability of mean spore length and width is usually
smaller, and that of mean @ considerably smaller at the popula-
tion level than at the individual level,
The variability of @ is much smaller with subglobose spores
than with cylindrical ones.
Spore size and spore form are characters widely used in
the taxonomy of fungi for distinguishing closely related
species. However, these characters have a statistical mean-
ing but are frequently used uncritically without taking
into consideration the extent of their variability. In our
earlier paper (E. Parmasto & I. Parmasto, 1982) we asserted
that the variability of these characters in the Aphyllopho-
rales is complicated and includes several components.
The coefficient of variation (or variability) y= ol
has been evaluated by Simpson, Roe & Lewontin (1960) as the
most convenient measure of variability. It has been widely
used in zoology and botany (including algology) but rarely
in mycology and probably not until now for the Aphyllopho-
rales.
This paper deals with the variability of mean spore
length, width and spore form index @ (ratio of mean length
to mean width) of five Phellinus species in 11 local popu-
lations. For comparison, the variability of Polyporus rhi-
zophtlus in one local population was also studied.
MATERIALS AND METHODS
The following species were studied: Phelltnus aint
(Bond.) Parm. (Ph. tgntartus (Fr.) Quél. f. alnz Bond.),
Pho Harttgt. (All. & Schn.) Pat., Ph. htppophaecola HL Jahn,
244
Phellinus igntartus (Fr.) Quel. s. str., Ph. robyustue
(P. Karst.) Bourd. & Galz., and Polyporus rhizophilus Pat.
Spore samples were collected as spore prints shed dur-
ing 24 hours from basidiocarps under natural conditions,
simultaneously from all the basidiocarps in each popula-
tion studied, except in two cases (P. rhtzophilus and
Ph. htppophaécola from Sweden), where spores of herbarized
basidiocarps collected within about one hour were measured.
From every sample 30 spores were measured in 2 % KOH
solution using an eyepiece micrometer at a magnification
of x 700 under the microscope MBM-6. For every sample the
mean values of spore length, spore width and @, and the
coefficient of variability Va of these data were calcu-
lated. Similar data were thereafter calculated for each
population, the coefficient of variation of sample means
within a population being denoted V oop" Details of the
samplings are given in Table l.
RESULTS are. summarized jin Table 2.
DISCUSSION
The number of specimens (samples) in the populations
studied was rather small (from 6 to 17) but the results
are similar enough to draw some conclusions, and Simpson,
Roe & Lewontin (1960) also used some rather small samples
(N = 8; 10) among their examples.
ee eT eee
The value of the variation coefficient VY was usually
4-10 for spore length anc width, and 3-8 for @ in the
Phellinus species. The mean value of V for @ (4.77) was
somewhat smaller than the ones for spore length (6.43)
and spore width (6.30); this is obviously connected with
the subglobose spore form. The mean value of VY for @ was
higher in Polyporus rhtzophtlus (10.3), which has cylin-
drical spores.
Variability of the means of specimens within one popula-
ELODs.i d nag” cull SON lade jhe AWee, 3 eee
The value of V was usually 2-7 for spore length and
width, and 1-2.7 for @ in the Phellinus species. The mean
value of VY for @ is much higher in Polyporus rhtizophilus
(5.7), which has cylindrical spores. Compared with the
variability of the spores of one basidiocarp, that of in-
dividual basidiocarps of the same population is usually
somewhat smaller for the mean spore length and width, but
always and remarkably smaller for Q.
Extent of variability in the fungi studied.
According to Rokickij (1973), the value of V ranges
mostly between 5 and 10 in homogeneous biological material.
Simpson, Roe & Lewontin (1960) as well as Mayr (1969)
245
asserted that the great majority of V’s for the linear di-
mensions of the anatomical elements of mammals lie between
4 and 10. Blue algae show similar or a little greater
variability (Kondratyeva, 1980). Our data fall within the
Samewcange; Obviously, the extent. of variab il <=
ipeyenCrom es se het iva Leia d Stic characters ds
Seiaueielyaer Sh pemos ts Org ah, items.
Variability between different populations of the same
species.
The comparision of data which may seem to be possible
for the Phellinus alnt and Ph. hippophaécola populations
has been avoided due to the fact that they were sampled
at different times. It has been demonstrated previously
(rarmasto: & E. Parmasto,. 1977), that spore size of
Phelltnus species may vary considerably within the same
specimens curing the sporulation period. Obviously, com-
parison of different populations in this group of fungi
yields reliable results only when the samples (from all
populations!) have been collected simultaneously, or in
sufficient numbers at quite different times and/or in the
case of basidiocarps at different stages of their devel-
opment. Even simultaneous collection of samples may only
yield suitable material when the collecting areas are not
too far from each other thus avoiding differences in
weather conditions.
Sometimes it is necessary to compare two specimens
(usually types) to establish their possible conspecific-
ity. Having measured a sufficient number of spores, it is
possible to use Student's t-test for establishing the sig-
nificance of difference between their mean measurements.
To evaluate such comparisions we studied two sets of
spore samples from Ph. tgntartus and Ph. htppophaécola
(from Sweden). Both sets were characterized by relatively
small variability of spores both of specimen and local
population levels.
Using the ¢-test in Ph. tgntarius all (136) possible
combinations (pairs) of specimens were compared and it
was established that the difference in the mean spore
Tength was highly significant (at P < 0.001) in 26 cases
animetqnitgrcant, (at .0.00l2< P< 0.01) ini 2) cases, Similar
results were obtained for Ph. htppophaécola: of 66 poss-
ible combinations (pairs) of specimens the difference was
highly significant in 20 cases and significant in 10 cases.
These results were obtained using specimens of one
local population characterized by a rather small coef-
ficient of variability and. collected at the same time.
Even in such "ideal" cases the comparision of two ran-
domly selected specimens using the t-test does not give
any useful information for the establishment of their spe-
cies identity or ditference. A biometrical comparison is
only reasonable between two populations, or between a
population and a single specimen.
246
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248
ACKNOWLEDGEMENTS
The authors are grateful to P.K. Mikhalevich, Esq.i,
and A.M. Zhukov, Cand. Biol. Sc. for the spore samples
from Byelorussia and Altai; to Dr. A.D.M. Rayner, Dr.
He Clémengon anc. a. Mols, Cand, Math. Sc. force: Lumcaiiy,
reviewing the manuscript, and to Asst. Prof. L. Kivimagi
for linguistic help.
LITERATURE CITED
Kondratyeva, N.V. (Edit.) 1980. Vnutrividovaya morfologicheskaya iz-
menchivost' sinezelenykh vodorosley (Cyanophyta). Kiev. (Intra-
specific morphological variability of blue algae. In Russian.)
Mayr, E. Principles of systematic zoology.
Parmasto, E. & Parmasto, I, 1982. Variability of spores of different
basidiocarps growing on the same mycelium in the Aphyllophorales.
Mycotaxon 16: 141-151.
Parmasto, I. & Parmasto, E. 1977. Variation in the dimensions of
spores of Phellinus sp. sp. during the discharge period. In: Eko-
logicheskie osobennosti nizshikh rasteniy Sovetskoy Pribaltiki.
Vilnius, p. 183-184. (In Russian, with an English summary.)
Rokickiy, P.F. 1973. Biologicheskaya statistika. Izdanie 3-e. Minsk.
(Biological statistics. In Russian.)
Simpsons, G.G., Roe, A, “~& Lewontin, R.C. 1960," Quantitative zool=
ogy. New York, Burlingame,
MYCOTAXON
MOE ALS No Viey “pp i249 -261 October-December 1982
NEW SPECIES GF FUNGILEROM THE
YUCATAN PENINSULA * **
by
Gast6n Guzman
Departamento de Bot&nica
Escuela Nacional de Ciencias Biolégicas, I.P.N.
Apartado Postal 26-378
México 16, D.F.
SUMMARY
Eight new species of fungi from the Yucatan Peninsula (Mexico) are
described. These are HYPHOMYCETES: Rhinotrichum mexicanum;
AGARICALES: Lepista singeri, Melanoleuca tropicalis, Amanita
ee rs er re SE
silvatica, A. yucatanensis. A. dunicola and Inocybe tropicalis, and
MY MENOGASTRALES: Octavianina cigroensis. All of them are
from the tropical rain forests, except Amanita dunicola which grows
in the dunes in the north of the peninsula where it is associated with
Coccoloba uvifera.
INTRODUCTION
During an extensive mycological exploration in the tropical
rain forests of Quintana Roo, and parts of Yucatan and Campeche
States, in November 1981, the author, in collaboration with the
Biologist Armando Lopez from Xalapa, Ver., found numerous in-
teresting fungi, lichens, and Myxomycetes, many of them newly
recorded in Mexico, of which eight species are described here.
Other (Lepiotaceae fungi) in Guzman Davalos and Guzman (1982).
All herbarium material is deposited in the following three
herbaria Mexican Institutes: Escuela Nacional de Ciencias Biolégi-
cas (ENCB) Mexico City, Instituto Nacional de Investigacién sobre
Recursos Bidticos (INIREB), Xalapa, Veracruz and Centro de Inves-
tigaciones de Quintana Roo (CIQRO), Puerto Morelos, Quintana Roo.
The latter has been provisionally deposited in MeridajYucatan ,
* Financed in part by CONACYT-BID, CIQRO and INIREB at
Mexico.
** Presented in the I National Congress of Mycology, at Xalapa
(Mexico).
250
The microscopic study .vas made from section mounted in KOH
(5%) or Melzer's solution. The colors of the spore, hyphae
and cystidia are described from mounts in KOH solution.
ACKNOWL EDG EMENTS
The author hereby expresses his gratitude to Dr. Alfredo Carrea-
ga, CIQRO Director, for his valuable help during the explorations
as well as to Dr. Arturo Gémez Pompa, INIREB Director and
M. enc. Alicia Barcena, Head of INIREB in Yucatan, for their
assistance to the program of the fungus flora of Yucatan Penin-
sula. The author also appreciates the good microscopic work and
herbarian work made _ by his daugher Laura Guzm&n DAvalos
and his assistants, Biol. S. Chac6n, Victor Hugo Ramirez,
Elsa Avilés Mercado and Tom&as Millan. Doctors A.L.Welden
from Tulane University and H.S. Thiers from San Francisco
State College, are also recognized by their interest and great
help in reviewing this article.
HYPHOMYCETES
Rhinotrichum mexicanum Guzman, sp. nov. Figs. ts
Mycelium flocci caespitosi, septati, brunneus ferrugineus,
hyphae pallide brunneae 6-12 um latae. Conidiophoris 11-25 X<
7-9 um, globosis vel ventricosis dein clavatus. Sterigmatis
numerosus breviculus. Conidiis 10-13 um Llatae, globosis,
hyalinae, tenuis verrucosis. Ad ligna vel corticem, subsilva
tropicalis. Quintana Roo prope, Coba-Nueva Xcan, Ldépez 1922
CTypus ENCB).
Mycelium ferrugineus brown, cottony, forming a loose
cushion-like mass on the substratum. Hyphae pale brown, septate,
intricately interwoven, 6-12 um diam. Fertile branches erec or
suberect. Conidiophores globose to ventricose or clavate, some-
times mucronate; with numerous short sterigmata over the surface,
11-25 X 7-9 pm; sterigmata not more than 3 um long. Frequently
the spores are borne on numerous sterigmata on cells below of
the conidiophore. Spores hyaline, globose, thick walled, finely
rough, 10-13 um diam.
Habitat. On bark in an open (disturbed) tropical rain forest.
Specimens examined. MEXICO, QUINTANA ROO, Coba to
Nueva Xcan road, near the crossing to Sajaca-chen, Ldépez 1922
(Type, ENCB).
Discussion. This fungus is close to R. subalutaceum Peck,
from NE of United States, but that species has spores 8-10ju m
diam., conidiophores which are not globose and mycelium that is
Figs. 1-18.- 1-3: Rhinotrichum, mexicanum, 1: Hyphae and
conidiophores, 2: Conidiophores, 3: Spores.- 4-9: Lepista
singeri, 4: Spores, 5: Basidium, 7: Trama,y S2sStructure of
the pileus, 9: Carpophores.— 10-14: Melanoleuca tropicalis,
10: Spores in KOH, ll: Spores in Melzer solution, 12: Basidia,
13: Cheilocystidia, 14: Structure of the pileus.- 15-18: Amanita
Silvatica, 15: Carpophores, 16: Elements of the volva on the
pileus, 17: Basidia, 18: Spores.
fais
yellowish brown. Itis also close to R. subferruginosum Sumstine
in the color of the mycelium and tropical habitat (Jamaica forests),
but that species has smooth spores (Sumstine, 1911). It is probable
that R. mexicanum has a wide distribution throughout the tropical
forests of Mexico (the author has observed this kind mycelium on
bark in forests in Veracruz State). Balfour-Browne (1952) des-—
cribed R. domesticum Balfour-Browne and R. Lanosum (Cooke)
Cooke as growing on walls in England, both with smooth spores.
AGARICAL ES
Tricholomataceae
Lepista singeri Guzman, sp. nov. Figs. 4-9
Pileo 50-80 mm lato, convexo vel applanato dein subdepresso,
levi, haud hygrophano, brunneolo griseus vel brunneolo alutaceus.
Lamellae subdecurrentibus vel subadnatis, subroseus. Stipite
45-55 X 10-15 mm, centralis vel subcentralis, albidus. Carne alba,
inmutabili, odore et sapore gratus vel fortis aromaticus. Sporis
5.4-7 X 3.5-4.5 um, hyalinae, globosis or ellipsoides, subtiliter
echinatis, haud amyloideis. Haud cystideis. WHyphis fibuligeris.
Quintana Roo, Puerto Morelos Prope, silva tropicalis, Guzman
21010: Giypus;- ENGB).
Pileus 50-80 mm diam., convex to plane or subconcave or
subinfundibuliform, smooth, sometimes with radical fibrills or
subrimose, dry, brownish gray or brownish leather color, with
some metalic shine, non hygrophanous. Lamellae subdecurrent
or subadnate , pale rose, more or less distant. Stipe 45-55 <
10-15 mm, central or excentric, equal or slightly tapering, bulbous,
whitish, fibrillose, smooth but somewhat floccose above, solid
to somewhat hollow, subcartilagineus, with white short rhizo-
morphs. Context white; taste and odor pleasant; dried specimens
with strong aromatic odor. KOH stains the pileus reddish brown.
Spores 5.4-7 X 3.5-4.5 yum, hyaline, globose or ellipsoid,
thin walled, finely echinulate, inamyloid. Basidia 28-37 X 8-9QumMm,
tetrasporic, hyaline, ventricose-fusoid. Pleurocystidia and
cheilocystidia absent. Trama regular, with hyaline hyphae
4.5-11 (-15)yim diam., thin walled, with numerous clamp connections.
Pileus surface with appressed hyaline hyphae, 3-6Um diam., some
with irregular incrusted brown pigment.
Habitat. Solitary or gregarious on humus, in the tropical
rain forests, below several species of trees, e.g., Achras zapota
(as ectomycorrhizic ?).
Specimens examined. MEXICO, QUINTANA ROO, near the
road from Puerto Morelos to Tulum, close to the crossing to Va-
llarta, footpath to the sea, Guzman 21010 (Type ENCB); Guzman
21018; GuzmA&n 21032; L6pez 1875. 10 km. from Tomas Garrido,
km 77 on road from Chetumal to Escarcega, Guzman 20883.
Discussion. This species is close to L. glabella (Speg.)
Sing. Known from Argentina, Paraguay, Brazil and Florida in sub-
tropical forests, but differs in the absence of an aqueous zone in
the gills close to the stipe, as well as in the presence of a brown
aqueous zone in the context above the gills, in the absence of an
odor and in the size of the spores (4.8-6.2 X 3.3-4.2 mm), accord-
ing Singer &Digilio (1951). None of the species of Lepista described
by Pegler (1977) from Africa agree with L. singeri. This is first
record of a Lepista in the tropical rain forests of Mexico. The
specimen 21018 was identified tentatively as L. singeri because it
is a young carpophore without spores, and with the pileus surface
formed from hyaline to somewhat incrusted more or less repent
hyphae 8-6um diam. The context has the same odor and the KOH
also stains the pileus brown reddish.
Melanoleuca tropicalis Guzm&n, sp. nov. Figs.l0-14
Pileo 15-35 mm lato, convexo vel subapplanato, glabro,
brunneolus griseus vel griseus plumbeus. Lamellis subadnatis,
albidis vel pallide cremeis. Stipite 20-40 X 1-2 mm, haud bubose,
albido, glabro. Carne albidae, odore nullo. Sporis (5.2-) 6-7.5
X 4.5-6 (-6.7) UM, globosis vel ellipsoideis, verrucosis, fortiter
amyloideis. Pleurocystidiis analogus cheilocystidiis, rarus.
Cheilocystidiis 31.5-55.5 &X 4.5-9 um, numerosus, obclavatis vel
lanceolatis, hyalinis, parte centralis septatis, ad apicem
acutissimis, crystallis hyalinis ad apicem incrustatis. Haud
fibulis. Ad terram in silva tropicalis. Quintana Roo, Puerto More-
los Prope, Santa Matilde, GuzmAn 21430 (Typus ENCB).
Pileus 15-35 mm diam., convex to somwthat plane or sub-
concave, glabrous, dry grayish brown to grayish lead color.
Lamellae subadnate, white or whitish to pale yellowish. Stipe
20-40 X 1-2 mm equal, white to whitish, glabrous. Context
whitish , odorless.
Spores (8.2-) 6-7.5 X 4.5-6 (-6.7) Um, globose or subellip-—
soid, hyaline, verrucose, thin walled, strongly amyloid, with a more
or less distinct plage on one side close to the apiculus. Basidia
16.5-28.5 X 9-10.5 um, hyaline, tetrasporic ventricose. Pleuro-
cystidia scarce, like the cheilocystidia but without apical crystals.
Cheilocystidia 31.5-55.5 X 4.5-9 uum, numerous, obclavate or
lanceolate, with a transver septum in the middle part and wth
characteristic hyaline apical crystals. Trama of more or less
interwoven hyaline hyphae. Pileus surface with appressed or sub-
appressed hyphae, with brownish gray content, 3-9 Um diam.
Clamp connections absent.
Habitat. Solitary on humus in the tropical rain forests.
Specimens examined. MEXICO, QUINTANA ROO, Puerto
Morelos to Vallarta road, 1 km. from the crossing to Tulum, Santa
Matilde, Guzm&n 21430 (Type ENCB). Puerto Morelos to Tulum,
close to the crossing to Vallarta, footpath to the sea, GuzmAn
21079.
Discussion. This species is close to M. brevipes (Bull. ex
Fr.) Pat. because of the cheilocystidia, but differs in the size of
the spores (8-10.5 X 4.5-6uUmM sensu Pegler, 1977; 7-8.5 X
4.5-6 um sensu Bon et al., 1973) and in the long stipe (short in
M. brevipes). Itis also close to M. pseudoluscina Bon but that
species has spores (6-) 7-8.5 (-9.5) X (5-) 6-7.5 (-8) Um (Bon,
1980). M. tropicalis is also close to M. tucumanensis Sing. from
Argentina (Singer &Digilio, 1951), but differs in the size of the
spores (7.5-10.3 X 6.2-7.5um) and in the basidia (GI1-35 X 8-9.7
Jum). Previous to this report, Melanoleuca was only known in
Mexico in temperate forests (Mendiola, 1974).
AMANITAC EAE
Amanita silvatica Guzm&n, sp. nov. Figs. 15-19
Pileo 20-35 mm lato, convexo dein applanate, subviscidus,
centro umbonatus, margin sulcatus striatus, aurantio luteus,
margin albidusy, fragmentum volvae tenuis, luteus, forma irregu-
lariter. Lamellae liberae, albae. Stipite 35-45 X 3-7 mm, albus,
exannulatus. Volva subsacciformis, alba, friabilis, marginis
aurantio luteus et fimbriate. Carne alba. Sporis (4.9-) 6.7-9
(-9.7) X (4.9-) 6-7.5 (-8.2) ppm, globosis vel subglobose, haud
amyloideis. Celluis volva 18.7-49.5 X 7.5-30 um, globosis,
fusiformis vel subellipsoideiss Fragmenta volva celluis pileo
(Q-) 25.5-43.5 (—45) X C6—-)16.5—-26.2(--30) sim, globosis vel sub-—
ellipsoideis et subfusiformis. Fibule absentes. Ad terram in
silva tropicalis. Quintana Roo, Puerto Morelos prope, Lépez
1847 (CTypus, ENCB).
Pileus 20-35 mm diam., convex to plane, smooth to sulcate-
striate at the margin, subviscid, yellowish orange to pale yellow
toward the margin, glabrous or with scattered, irregularly
shaped, yellowish, rather thick, floccose-membranous flat
patches from the volva. Lamellae free, white to cream. Stipe
35-45 X 3-7 mm, equal or attenuated upward, base globose or
subglobose, white to whitish cream, somewhat floccose be low.
Annulus absent. Volva as an adhering sac, with the upper part
free, with the margin fimbriated or lacerated, white to orange
yellowish on the margin. Context white, odor not distinctive.
Los
Spores (4.9-)6.7-9(-9.7) X (4.9-)6-7 .5(-8.2) um, globose
to subglobose, with a short apiculus, hyaline, inamyloid. Basidia
19.5-22.5 X 9-9.7 Im, tetrasporic, hyaline, ventricose. Cells
of the margin of the volva 18.7-49.5 X 7.5-30 tim, globose, fusi-
form or subellipsoid, hyaline, thin walled, borne on hyphae
1.5-13.5 pam broad. Cells of the volva on the pileus globose, sub-
fusiform or subellipsoid, (9-)25.5-43.5(-45) X (6-)16.5-26. 2(-30)
Jum, hyaline, thin walled, borne on hyphae 3-12 um broad. Clamp
connections absent.
Habitat. Subgregarious on soil in a tropical rain forest
(as ectomycorrhizic 7).
Specimens examined. MEXICO, QUINTANA ROO, near the
road from Puerto Morelos to Tulum, close to the crossing to Va-
llarta, footpath to the sea, Ldépez 1847 (Type, ENCB).
Discussion. A. silvatica is close to A. elata (Mass.) Corner
& Bas, which is Known only from Singapore and differs in the size
of the cells of the volva at the stipe (40-60 X 25-40 prn), presence
of annulus, in the difference in the color of the pileus (pale dingy
ochraceous buff or dingy buff with sulphur yellow tones) and in the
more narrow volva at the bulb without fimbriate edges (Corner &
Bas, 1962).
Amanita yucatanensis Guzman, sp. nov. Frgs. 20-26
Pileo circa 25 mm lato, convexo dein subapplanato, margin
sulcatus, siccus, albus vel subflavidus, fragmentum volvae albidus,
tenuis, forma irregulariter. Lamellae subliberae, albae. Stipite
circa 12 X5 mm, albus, squamosus, exannulatus. Volva sacci-
roOrinis, alba, tenera. Sporis (7.5—)9-12C¢12.7) xX @.1-)s.2-9
(-10.5) Um, globosis, haud amyloideis. Celluis volva subglobosis,
30-52.5 X 12-21 um, hyalinae. Fibule absentes. Ad terram in
silva tropicalis. Quintana Roo, Puerto Morelos prope, Guzman
21044 (Typus, ENCB).
Pileus about 25 mm diam., convex to somewhat plane,
smooth to striate at the margin, dry, white to pale yellowish ,
glabrous or with some scattered irregularly shaped, whitish,
thin, flat patches of the volva. Lamellae subfree, white to
whitish rose, with fimbriate edges. Stipe aboutl2 x 5 mm,
very attenuated upward, floccose-scaly to pruinose above,
without annulus. Volva more or less high and wide, as a mem-
branous sac, white and delicate. Flesh white, odorless.
Spores (7 .5-)9-12(-12.7) <(7.1-)8..2-9¢ -10.5) pm, globose
with a short apiculus, hyaline, inamyloid. Basidia 42-54 <
10.5-13.5 jum, tetrasporic, but some mono or bisporic, hyaline,
256
ventricose, with a middle constriction. Cystidia none. Trama
with elongated to broad cells, hyaline. Volva with subglobose hya~
line cells, 30-52.5 X 12-2lum, borne on narrow hyphae 2.2-9 pum
broad, many of them with grayish irregularly incrusted pigment
on the walls. Epicutis formed by more or less repent hyaline
hyphae, 3-7.5 pam broad. Hypodermium with subglobose hyaline
elements, 9-19.5 pum broad. Clamp connections absent.
Habitat. Solitary on soil in a tropical rain forest.
Specimens examined. MEXICO, QUINTANA ROO, Road to
Vallarta, near the crossing from Puerto Morelos to Tulum road,
Guzman 21044 (Type, ENCB).
Discussion. This species is very close to Amanita species
No. 4 of Corner &Bas (1962), Known only from Singapore, but
differs in the spores (Q9-11.5 X 7-8.7 pum). The white basidiocarp
seems similar to A. praegraveolens (Murr.) Sing., also a tropical
species, but that fungus has scaly floccose, poorly formed volva
and amyloid spores (Guzm&n 1975).
Amanita dunicola Guzman, sp. nov. Figs. 24-28
Pileo 15-40 mm lato, convexo dein subcampanulatus vel
subapplanato, subviscidus, sulcatus-striatus, subbrunneolus vel
alutaceous, fragmentum volva albus tenuis, forma irregulariter.
Lamellis liberae, albidus roseus. Stipite 30-80 XK 3-7 mm, albus
vel inaequaliter brunneolus, exannulatus. Volva sacciformis,
alba, tenera. Sporis @.9-)l1-12.5(-13.5) X 8-9 xsm, subglobosis,
non amyloideis. Celluis volva globosis 17-38 um. Fibule in the
basidia. Ad arena contiguus ad oceanus. Yucatan prope, circa
Ixil, Progreso-Telchac, Guzm&n 21235 (Typus, ENCB).
Pileus 15-40 mm diam., convex to subcampanulate or some-
What plane, smooth to sulcate-striate at the margin, subviscid,
pale brownish to leather brown, with white, irregularly flat
patches from the volva. Lamellae free, whitish rose to rose brown,
with white fimbriate edges. Stipe 30-80 X 3-7 mm, equal or some-
What attenuate at the base, smooth but floccose under the lens,
white to irregularly brownish. Annulus absent. Volva as a mem-
branous sac, white and delicate, more or less permanent in the
adult. Context white, odor not distinctive (but in dry specimens
very pleasant, like bread); taste pleasant.
Spores (9.9-)l1-12.5(-13.5) X 8-911m, subglobose with a
short apiculus, hyaline, inamyloid. Basidia 50-70 X 13-16 um,
tetrasporic, hyaline, clavate; sterigmata up to €um long.
Cystidia absent. Trama with hyaline elongated hyphae, 3-6 um
broad, some anastomoses. Cuticle of the pileus subgelatinous,
formed by hyaline to brownish appressed hyphae up to 5 pum
Figs. 19-34.- 19: Amanita silvatica, elements of the volva
on the stipe.-— 20-23: Amanita yucatanensis, 20: Spores, 2l:
Basidia, 22: Structure of the pileus, 238: Elements of the volva.-—
24-28: Amanita dunicola, 24: Carpophores, 25: Spores, 26:
Young basidium with a clamp connection, 27: Basidium, 28: Ele-
ments of the volva at the stipe.- 29-32: Inocybe tropicalis, 29:
Carpohore, 30: Spores, Sl: Basidium, 32: Cheilocystidia.-—
33-34; Octavianina cigroensis, 33: Spores, 34: Carpophores.
256
broad. Elements of the volva globose, hyaline, 17-38um broad,
thin to thick walled, borne on hyphae 3.3-6Hm broad. Clamp
connections observed only at the base of the basidia.
Habitat. Gregarious, rarely solitary, in sand below
Coccoloba uvifera, in dunes close to the sea. It seems to be an
ectomycorrhizic fungus.
Specimens examined, MEXICO, YUCATAN, Municipio de
Ixil, Progreso to Telchac road, Guzm&n 212385 (Type, ENCB).
Discussion. This species is close to A. antillana Dennis
from Trinidad, but differs in the friable volva, in the scaly-floc-
cose stipe, broader spores (7.5-11.541m), glabrous and olive
brown or grayish pileus, and in the habitat (ad terram in silvis)
(Dennis, 1952, 1970). Itis also close to Amanitopsis floridana
Murr. from Florida, because of color of its pileus and non friable
volva, but it has narrower (6-7 sum) spores and grows on soil close
to pines (Murrill, 1949). Amanita species No. 5 of Corner &Bas
(962) from Singapore is also close, but differs in its glabrous
pileus and broader spores (9.5-l1 um).
CORTINARIAC EAE
Inocybe tropicalis Guzm&n, sp. nov. Figs. 29-32
Pileo 20-25 mm lato, conic, bruneus aurantius, vel
brunneolus, fortiter rimosus. Lamellis brunneus aurantiacus
cum albidus fimbriate marginis. Stipite 16-18 X 3-5 mm, albidus
vel rufobrunneus. Carne albidus, odor nullus. Sporis (8.8-)
12-13(-17 .6) X 5.5-6.6(-7.7)um, subellipsoideis vel oblongatus-—
ellipsoideis. Plecrocystidiis absens. Cheilocystidiis 44-55 x
12-27.5um, hyalinea, subclavate vel subturbinate, sine crustare.
Epicutis cum parallelis vel suberigo hyphae, 7-15 (—20) um latus.
Hyphae fibululigeris. Ad terram, solitario, in silva tropicalis.
Quintana Roo, Tomas Garrido prope, Guzman 20927 (Typus,
ENCB).
Pileus 20-25 mm diam., conic, brownish orange to pale
brown, strongly radially fibrillose and rimose, revealing the pale
context. Lamellae subadnate, brownish orange with white fimbria-
te edges. Stipe 16-18 K 3-5 mm, uniform, whitish to yellowish,
finally brownish red, smooth, somewhat cortinate in the middle
portion. Context whitish, odorless,
Spores (8.8-)L2-13(-17 .6) X 5.5-6+6(-7.7) jum, subellipsoid or
oblong-elliptical, brownish mustard, with a narrow germ pore.
Basidia 35-40 X ll-15pim, tetrasporic, hyaline, ventricose-clavate,
with sterigmata up to 2um thick. Pleurocystidia absent. Cheilo-
cystidia 44-55 X 12-27.5 Mm, numerous, hyaline, subclavate or
subturbinate, without incrustations. Subhymenium pale brown-
ish mustard. Trama regular, hyaline. Pileus surface with
prostrated to somewhat erect incrusted brownish orange pigment—
ed hyphae, 7-1L5(-—20)H¥m broad. Clamp connections observed in
several hyphae, even at the base of basidia and cheilocystidia.
Habitat. Solitary on soil in tropical rain forest (ectomy-
COrEhiZic 2).
Specimens examined. Mexico, Quintana Roo, 10 km N of
Tomas Garrido (South of Quintana Roo), Guzm&n 20927 (Type
ENCB).
Discussion. I. tropicalis is somewhat close to I. jalapensis
(Murr.) Sing. but differs in the presence of pleurocystidia (55-80
X 16-20 f’m) in that species as well as in the size of the spores
(7 .7-9.8 X 4.2-5.7p mM), and in the subtropical distribution.
Murrill described that species from Xalapa, Ver. and Dennis re-
ported it from Colombia (Singer, 1957; Dennis, 1970), in both
cases itis ectomycorrhizic and associated with Quercus. Itis also
close to I. jamaicensis Murr. but that species has rugose spores
8-9 X 5U¥m (Murrill, 1912). The only species of Inocybe reported
by Pegler (1977) from Africa is I. lLanuginella (Schroet.) Konr. &
Maubl. from a Pinus forest in Tanzania, which has pleurocystidia.
HYMENOGASTRALES
Octavianina cigroensis Guzman, sp. nov. Figs. 33-34
Carpophoris 5-10 mm Lato, globosis vel subglobosis. Peridio
luteus vel spadiceus. Gleba albidus vel spadiceus. Sporis (8.2-)
9-12(-13.5) X (7.5-)9-9.7(-10.5)um, globosis, pseudoamyloideis,
verrucosis subreticulatis. Cystidiis nullis. Hypogaeum in silva
tropicalis. Quintana Roo prope, Carrillo Puerto-Vijia Chico,
Ldépez 1759 (Typus, ENCB).
Gastrocarp globose or subglobose, 5-lL0 mm diam. Peridium
glabrous, with some innate veins especially below, thin, yellow,
fading to leather brown. Gleba chambered, whitish to pale brown-
ish,columella and sterile base absent.
Spores (8.2-)9-12(-13.5) X(7.5-)9-9.7(-10.5) pim, globose,
hyaline, verrucose, subreticulate, pseudoamyloid, with a short
but conspicuous appendage. Cystidia none. Peridium with irregu-
larly elongated hyaline to pale brownish hyphae, without clamp con-
nections.
Habitat. Hypogeous in humus of a tropical rain forest.
260
Specimens examined. MEXICO, QUINTANA ROO, 20 kms
from Felipe Carrillo Puerto, road to Vijia Chico, Lépez 1759
(Type, ENCE).
Discussion. O. tuberculata (Hesse) 0. Kuntze and 0. laevis
(Hesse) O. Kuntze are the only species somewhat close to O.
cigroensis following Singer & Smith (1960) key's, but they differ
in diameter and ornamentation (broad cones) of the spores (12-16
Mm diam. in the first, and 13-18(—20) pm diam. in the second).
Both species are only known from Germany. Apparently 0.
ciqroensis is the first record of the genus in the tropics, at least
in Mexico.
The name of the species refers to CIQRO Institution (Centro
de Investigaciones de Quintana Roo) in recognition to the support
given to this research project through its Director, Dr. Alfredo
Carreaga.
LITERATURE Cit ED
Balfour-—Browne, F.L., 1952. Three moulds from damp walls.
Trans. Brit. Myc. Soc. 35:278-278.
Bon, M., 1980. Agaricales rares ou nouvelles de la Zone Mariti-
me Picarde, 5. Bull, Soc. Myc... 90:S50=174.
Bon,/M., G..Ghevassut, P.«Bertea, CG. Eecot and J. CL. Rousse,
1973. Agaricales de la Region Languedoc-Cevennes. Doc.
Corner, E. J. H. and C. Bas, 1962. The genus Amanita in
Singapore and Malaya. Persoonia 2:241-304.
Dennis, R. W., 1952. Lepiota and allied genera in Trinidad,
British West Indies. Kew Bull. 7:459-499.
21970. Fungus flora of Venezuela and adjacent countries.
Kew Bull. Addit. Ser. III and Cramer, Lehre.
Guzman, G., 1975. New and interesting species of Agaricales of
Mexico;.in Bigelow, H. E. andH. D. Thiers, STUDIES Gy
HIGHER FUNGI. Beih. Nova Hedwigia 51, Cramer, Vaduz.
Guzman-D4avalos, L. and G. Guzm&n, 1982. Nuevas especies de
Lepiota. Bol. Soc. Mex. Mic. 17 (in press).
261
Mendiola, G., 1974. Estudio taxonémico y datos ecolégicos de
algunas especies mexicanas de hongos de la Familia Tricho-
lomataceae (Agaricales). Escuela Nacional de Ciencias Bio-
légicas, I.P.N., México, D.F. (thesis).
Murrill, W. A., 1912. The Agaricaceae of Tropical North Ame—
rica. Mycologia 4:72-83.
21949. Three new fleshy fungi. Mycologia 41:490-491.
Pegler, D. N., 1977. A preliminary agaric flora of East Africa.
Kew Bull. Addit. Ser. VI and Her Majesty's Stat. Office,
London.
Singer, R., 1957. Fungi Mexicani, series Prima, Agaricales.
Sydowia 11:354-374.
» andA. P. L. Digilio, 1951. Prdédromo de la flora agaricina
argentina. Lilloa 25:5-46l1.
» and A. H. Smith, 1960. Studies on Secotiaceous fungi,
Sumstine, D.R., 191ll. Studies in North American Hyphomy-
cetes, I. The genera Rhinotrichum and Olpitrichum,
Mycologia 3:45-56 +3 pls.
MYCOTAXON
Vol xXViilw eNO leepp 267-264
October-December 1982
TWO NEW FAMILIES IN THE ASCOMYCOTINA
D. L. HAWKSWORTH
Commonwealth Agricultural Bureaux, Farnham Royal, Slough SL2 3BN, UK
and
M. A. SHERWOOD
P.O. Box 934, Marcola, Oregon 97454, USA
SUMMARY
Two new families are described in the Ascomycotina for
the first time. Ascodichaenaceae fam. nov. (Rhytismat-
ales) is introduced for Ascodichaena, Delpinoina, and
(?) Pseudophacidium. Odontotremataceae fam. nov. (Ost-
ropales) is described for Bryodiscus, Lethariicola,
Odontotrema, Odontura, Skyttea, Spilomela, Stromatoth-
ecia and Xerotrema.
In the course of our investigations on the systematics of the
Ascomycotina, it became clear that two groups of these fungi were
worthy of recognition in the rank of family. These two family names
are formally described here and their circumscription and affinities
discussed.
ASCODICHAENACEAR D.Hawksw. & Sherw. fam. nov.
Familia ad ordo Rhytismatales pertinens. Ascomata stromatica,
erumpescentia, aggregata, carbonacea, disciformia ad breve hyster-
iformia, longitudinaliter apperientia, excipulum vero desunt, stro-
mata ex cellulis pseudoparenchymatis polyedricis et verticalis.
Paraphyses numerosae, simplices, septatae. Asci cylindrici sed cE
avati ubi maturitati, tenue, cum apicibus late annulatis, probaliter
operculati, 4-8-spori. Ascosporae hyalinae, simplices.
Anamorphosis holoblasticae vel ignota.
Holotypus: Ascodichaena Butin.
Family belonging to the order Rhytismatales (syn. Phacidiales
auct.). Stromata erumpent in dense clusters, carbonaceous, true
excipulum absent, stromatal tissue orbicular to short-hysteriiform,
opening by a longitudinal slit, composed of vertically orientated
polyhedral pseudoparenchyma. Paraphyses numerous, simple, septate.
Asci at first cylindrical and sessile, becoming clavate or saccate
as the ascospores mature, thin-walled, I-, with a broad apical
annulus, possibly operculate, 4-8 spored. Ascospores large, colour-
less, unicellular, lacking a gelatinous sheath.
Anamorph coelomycetous, holoblastic (Polymorphum Chev.), or
unknown; spermatial phase sometimes present.
Habitat: On the bark of Fagaceae in temperate regions of the
Northern Hemisphere.
263
Genera: Ascodichaena Butin, Delpinoina Kuntze (syn. Henriquesia
Pass. & Thum.) and possibly Pseudophacidium P. Karsten.
Relationships: Butin (1977) summarized earlier treatments of
Ascodichaena, concluding that it was most closely allied to Pseudo-
phacidium and referring it to the family Phacidiaceae Fr. However,
neither Ascodichaena nor Pseudophacidium have an ascus with an amyloid
apical ring such as that seen in Phacidium Fr. This distinction, and
the presence of a holoblastic anamorph in Ascodichaena and a phialidic
one in Phacidium, indicates that the former is most appropriately
placed in the Rhytismatales. Further, the distinctive anamorph, the
stroma, and absence of sheaths on the ascospores, justifies the
placement of Ascodichaena in a separate family within that order.
The family name Dichaenaceae Fr. cannot be taken up for this
family as the type species of Dichaena Fr. is anamorphic (Hawksworth
& Punithalingan, 1973). Speer (1980) considered that Dichaena was a
teleomorphic generic name but his conclusions were based on a number
of misinterpretations of the Code which will be discussed by DLH in a
separate paper.
On the basis of ascus structure, the Phacidiaceae is most appropr-
iately referred to the Helotiales and has strong affinities with the
Dermateaceae Fr. In addition to Phacidium, the Phacidiaceae comprises
Lophophacidium Lagerberg, | Phacidiostroma Héhnel, and perhaps also
the genera Cryptomycina Héhnel, Nannfeldtia Petrak, Micraspis Darker
and Phacidina Hohnel.
ODONTOTREMATACEAE D.Hawksw. & Sherw. fam. nov.
Familia ad ordo Ostropales pertinens. Ascomata hemiangiocarpia,
apothecia, immersa vel erumpescentia, marginata, singularia vel in
stromatis aggregata; excipulum ex hyphis compositum, usque atrobrunnea
ad nigra, interdum carbonacea. Asci unitunicati, cylindrici, cum
apicibus crassis, plusminusve pori, cum iodo non reagens, 8- vel multi-
spori. Ascosporae ovoideae, sigmoideae, vel filiformiae, hyalinae,
septatae ad muriformiae.
Anamorphosis ignota.
Holotypus: Odontotrema Nyl.
Family belonging to the order Ostropales. Ascomata hemiangiocarpic,
apothecioid, immersed or erumpent, determinate, marginate, single or
immersed in a stroma; excipulum hyphal, usually dark coloured and in
some genera carbonaceous. Paraphyses simple or branched at the base,
septate. Asci cylindrical, sessile or nearly so, the lateral walls
thin and the apex thickened, I-, with or without an apical pore,
inoperculate, 8- or multispored. Ascospores ovoid, sigmoid, or fili-
form, hyaline, simple, transversely septate or muriforn.
Anamorph not definitely known (a pycnidial fungus was illustrated
by Grummann, 1969, associated with Lethariicola apothecia but its
connection with them is uncertain).
Habitat: Saprobic, chiefly on wood in xeric situations, or lichen-
icolous (and then usually parasymbiotic).
Genera: Bryodiscus Hein et al., Lethariicola Grumm., Odontotrema
Nyl., Odontura Clem., Skyttea Sherw. et al., Spilomela anne \Keissl.,
264
Stromatothecia D.Hawksw. & D.Shaw, and Xerotrema Sherw. & Coppins. It
is also probable that the lichen-forming genera Bryophagus Nitschke ex
Arnold and Ramonia Stizenb. should be referred to this family.
Relationships: This distinctive assemblage of genera was noted
by Sherwood et al. (1981) to almost certainly form a natural unit. No
new family name was introduced at that time as the application of
some other names, especially Triblidiaceae Rehm, was then uncertain.
The latter family includes Pseudographis Nyl. and Triblidium Rebent.,
Tryblidiopsis P. Karsten being more appropriately placed in the order
Rhytismatales. The Triblidiaceae differs from the Odontotremataceae
in that the covering stroma splits stellately or by a slit to expose
the hymenium, the presence of a distinct epithecium, and ascospores
which are either transversely septate with lenticular cells or
muriform, turning reddish-purple in iodine. This last reaction may
indicate some relationship to the Graphidales. The asci in the Tribl-
idiaceae have a strongly thickened apex which can be clearly seen to
be pierced by a pore; they are I- as is the case in the Odontotremat-
aceae and Graphidaceae.
ACKNOWLEDGEMENTS
We are grateful to numerous colleagues for stimulating discussions
on the classification of the taxa recognized here, but especially wish
to thank Frank Di Cosmo for his views on the Phacidiaceae. The work on
which this paper is based was undertaken whilst MAS was employed as a
Senior Research Assistant at the University of Liverpool on a Science
Research Council funded project for the preparation of a new checklist
of British ascomycetes.
REFERENCES
Butin, H. (1977) Taxonomy and morphology of Ascodichaena gen. et sp.
nov. Trans. Br. mycol. Soc. 69: 249-254.
Grummann, V.J. (1969) Alte und neue Halbflechten. Ein neuer Flechten-
parasit. Placynthium asperellum neu fur Mitteleuropa. Sydowia 22:
216-224,
Hawksworth, D.L. & Punithalingam, E. (1973) Typification and nomencl-
ature of Dichaena Fr., Heterographa Fée, Polymorphum Chev., Psil-
ospora Rabenh. and Psilosporina Died. Trans. Br. mycol. Soc. 60:
501=509..
Sherwood, M.A., Hawksworth, D.L. & Coppins, B.J. (1981) "1980" Skyttea,
a new genus of odontotremoid lichenicolous fungi. Trans. Br. mycol.
Soc. 75: 479-490.
Speer, £.0. (1980) Dichaena rugosa Fries and its imperfect states,
Psilospora faginea (Pers.) Rabenh. and Dichaenopsis notarisii Paoli.
Taxon 29: 641-644,
MYCOTAXON
WOMePN WE NO w st, apps 2652272 October-December 1982
Se ee ee cae Ea a Sak einen oleate ed le ce ara
]
CYLINDROCLADIUM SPATHIPHYLLI SP. NOV.
CALVIN L. SCHOULTIES, NABIH E. EL-GHOLL,
AND SALVATORE A. ALFIERI, JR.
Bureau of Plant Pathology
Divtston of Plant Industry
Flortda Department of Agrtculture & Consumer Services
Post Of fice Box 1269
Gatnesvtlle, FL 32602
ABSTRACT
Cylindrocladium spathiphylli is described as a new
species from Spathiphyllum sp. 'Clevelandii' in Florida.
INTRODUCTION
Beginning in late 1978 and continuing to the pre-
sent, several nurseries in Florida have incurred serious
economic losses from a Cylindrocladium root and foliar
disease of various species of Spathiphyllum. We have
heretofore designated the pathogen as Cylindrocladium
floridanum Sobers and Seymour (1967) in previous publi-
cations (Schoulties and El-Gholl, 1980a, 1980b, 1981).
However, the isolate described from Spathiphyllum is now
known to be different from other described species of
Cylindrocladium. The following presents the distinctive
morphological features of the fungal isolate from Spath-
iphyllum and provides the basis for establishing it as a
new taxon.
METHODS AND MATERIALS
The isolate of Cylindrocladium under study was
obtained from basal portions of petioles and from roots
medae ei bution No. 256, Bureau of Plant Pathology.
266
of Spathiphyllum sp. 'Clevelandii' which was being grown
at a commercial nursery in Apopka, Florida. The isolate
of C. floridanum was obtained for comparative study from
leaf spots on leatherleaf fern, Rumohra adiantiformis
(G. Forst.) Ching which was being grown in Pierson,
Florida.
Potato dextrose agar (PDA) was prepared from the
broth of 200 g freshly peeled, diced, and boiled Irish
potatoes supplemented with 20 g dextrose, 1 g KH,PO
and 18 g Difco bacto-agar, and made up to 1 liter with
deionized water.
Peanut stem water agar (PSWA) was prepared in the
following manner: cooled (48 C), autoclaved (121 C for
15 min) water agar was poured over dried, propylene
oxide-fumigated (Hansen and Synder, 1947) stem pieces of
peanut. Under careful handling, the stem piece floated
and became partially submerged upon solidification of
the agar.
To obtain monoconidial cultures, the method de-
scribed by Hansen and Smith (1932) was used.
Cultures of Cylindrocladium were grown at 25 C
under 12-hour periods of alternating light (fluorescent
light, Westinghouse F20T12/CW at an intensity of approx-
imately 2800 lux) and dark on either PDA or PSWA for 10
days.
Morphological features were detailed and measured
under oil immersion and were photographed (Fig. 1) with
Nomarski differential interference-contrast microscopy
using a Zeiss photomicroscope III. Spore measurements
represent 400 conidia.
TAXONOMY
The Cylindrocladium from Spathiphyllum sp. 'Cleve-
landii' differs from other species of Cylindrocladium in
the following characteristics. It has l-septate conidia
and a stipe terminating in a globose vesicle, differing
from Cylindrocladium ilicicola (Hawley) Boedijn and
Reitsma (1950), C. crotalariae (Loos) Bell & Sobers
(1966), and C. citri (Fawcett & Klotz) Boedijn and
Reitsma (1950) which have globose vesicles and 3-septate
conidia. It differs from other species of Cylindro-
cladium with globose vesicles and l-septate conidia in
having cylindrical conidia 45.0-101.0 x 5.0-7.0 um as
compared to 40-46 x 3-4 um for the curved conidia of C.
curvatum Boedijn and Reitsma (1950) and 30-56 x 3.5-5.4
267
Fig. 1. Cylindrocladium spathiphylli: A) Single
central stipe terminating in a globose vesicle, 560 X.
B) Conidiophore arising from young procumbent mycelial
cell, 560 X. C) One-septate conidia, 560 X. D) Conidi-
ophore arising from a sclerotial cell with a ruptured
cellular membrane (collarette - noted by arrow), 1400 X.
268
yum for the cylindrical conidia of C. floridanum Sobers &
Seymour (1967). We obtained the following conidial
measurements on an isolate of C. floridanum from leather-
leaf fern: (39.5-) 47.2 (-56.0) um long by (3.0-) 3.9
(-4.0) um wide. Further, C. floridanum is known to have
in addition to the central stipe (an extension of the
main axis of the conidiophore stalk) lateral stipes from
secondary and tertiary conidiophore branches (Morrison
and French, 1969), terminating in sphaeropedunculate
(Snell and Dick, 1971) vesicles with a somewhat flat-
tened top, whereas the Cylindrocladium from Spathi-
phyllum has but a single, central stipe per conidio-
phore, terminating in a larger globose, nonsphaeropedun-
culate vesicle, as well as having larger and wider
conidia. Descriptions were based on a monoconidial
isolate and were compared with original descriptions of
Cylindrocladium species, Matings among various isolates
from Spathiphyllum never produced the teleomorphic state
under the conditions studied, hence the species appears
to be heterothallic.
On the basis of the differences noted as outlined
in Table 1, this isolate is considered morphologically
distinct from previously described species in this genus
and we propose the following new species:
Cylindrocladium spathiphylli Schoulties, E1-Gholl
et Alfieri, sp. nov.
Aerium mycelium coloniae in agaro Solani tuberosi,
bene maturum, initio album et gossypinum, aetate varie
brunneum vel rufum. Conidiophora lateraliter portata
stipite suragente ex cellulis mycelii procumbentis vel
ex cellulis sclerotiorum quae cellulae membranum cell-
ulare ruptum, ut collare parvulum, ad basim stipitis
praebent. Conidiophorum ramificatio dichotoma. Rami
primarii hyalini, leves, non septati, aliquando uni-
septati,, (19%0-) 32.8-(-57.0). um x (4.0) 5.1 (-6.5 8a
rami secundarii vel metulae hyalini, leves, non septati,
(13.0-) 21.6 (-35.0) um x (4.0-) 4.6 (-6.0) um; rami
tertii, cum adsunt, hyalini, leves, non septati (18.0-)
20.3 (=2220) jm ‘x 4.0 um. ~Phialidese2-4) .cylLindratac,
fine apicis obtusae, vel aliquando doliiformes vel
reniformes, hyalinae, non septatae, (10.0-) 14.6 (-20.0)
um x (3.0-) 4.2 (-5.0) um. Stipites vel ex hospite vel
ex myceliis in cutura procumbentibus vel ex corporibus
sclerotiorum in hospite et in cultura recte surgentes,
269
“(ZL61T) Sateqos pue (g96T) BITYSPIAL,,
°(1961) Anoudss pue S19q0s_
* (eseraAe **o°* Tt) UsATS JON = ONG
°(6961) Youerq pue UoSTIIOWL,
aqeTnounped O20 [LOS ts | ue hoeor
umoux jueseid -oreeyds (0°S-)0°7(-0°€) (O°IS-)o'I7(-0'6z) © -opnesd epuTqoy
ae TNounped
umouyx Juesead -ozseyds = (g* #-) DN(-G*€) (0°SS-)ON(-0° 7€) 4278T0ST yoved
33 eTNounped 33eTOST
umouy Jueseid -oteeyds (0°7-)6°€(-0°€) (O° 99S) 2510-9. 6G) Uldey JeeTYAeST
93eTNounped
umouy jueseid -oreeyds = (€*¢-)9N(-0°7) (0°6S-) ,ON(-0°€€) prtelOst tazTuoo
WNUEPTIOTF °9
umouy Jou juesqe es0qgoTs (0°/-)0°9(-0°S) (0°101T-)€°08(-0'S¥) FrItAydryjzeds +5
ALVLS SHdILS AdVHS (url) HLGIM (ur) HLONAT Sa10adS
TWAXdS ‘IVYALVI ATOISHA SINAWAYNSVAW TVICINOO
“eTpTuod ojejdes suo sey osTe Jey WnuepTAOTJ uUNTpeTooapuTTAD UIIM
TrtAudtyjeds wntpepoorputpsé9 jo seanjzeey TeotTZoToydizow SAFIOUTISTP Jo uostazaedwoy °*T eTgqey],
270
septati, ad basim dilute brunnei, (7.0-) 7.8 (-9.0) um
lati, (111.0-) 140.8 (-253.0) um longi, ad apicem sensum
hyalini et angustiores (3.0-3.7 um lati), vesicula
hyalina, granulari, globosa, (9.0-) 12.5 (-15.0) um
diam. terminati. Conidia ex apice phialidum singulatim
ficta et in circulis vallo similibus et substantia
mucilaginosa contentis cumulata, hyalina, granularia,
cylindrata, levia, recta, utrisque finibus rotundata,
uniseptata, (45.0-) 80.3 (-101.0) um longa, (5.0-) 6.0
(-7.0) um lata.
Segregatum ex partibus basalibus petiolarum et ex
radicibus Spathiphylli sp. 'Clevelandii' in Apopka,
Florida. Depositum ut ATCC 44730 in American Type
Culture Collection, 12301 Parklawn Drive, Rockville,
Maryland 20852, U.S.A.
Cylindrocladium spathiphylli Schoulties, El-Gholl &
ALS Lerigsp. nov.
Aerial mycelium of colony on potato dextrose agar,
well developed, initially white and cottony, becoming
variously brown to reddish brown with age. Conidio-
phores are borne laterally on a stipe arising from cells
of procumbent mycelium or from cells of sclerotia that
produce a ruptured cellular membrane seen as a collar-
ette at the base of the stipe. Conidiophore branching
is dichotomous. Primary branches hyaline, smooth,
nonseptate, occasionally l-septate, (19.0-) 32.8 (-57.0)
um x (4.0-) 5.1 (-6.5) um; secondary branches or metulae,
hyaline, smooth, nonseptate, (13.0-) 21.6 (-35.0) um x
(4.0-) 4.6 (-6.0) um; tertiary branches when present,
hyaline, smooth, nonseptate (18.0-) 20.3 (-22.0) um x
4.0 um. Phialides 2-4, cylindrical with obtuse apical
end to occasionally doliiform or reniform, hyaline,
nonseptate, (10.0-) 14.6 (-20.0) um x (3.0-) 4.2 (-5.0)
um. Stipes arise at right angles from the host or from
procumbent mycelia in culture or from sclerotial bodies
on the host and in culture, septate, light brown at the
base (7.0-) 7.8 (-9.0) um wide, (111.0-) 140.8 (-253.0)
um long, becoming hyaline and narrower at the apex (3.0-
3.7 um wide), terminating in a hyaline, granular, glo-
bose vesicle (9.0-) 12.5 (-15.0) um diam. Conidia are
formed singly from the apex of the phialides and ac-
cumulate in palisade-like clusters held together by a
mucilaginous substance. They are hyaline, granular,
cylindric, smooth, straight, rounded at both ends, l-
Aig
septate (45.0-) 80.3 (-101.0) um long by (5.0-) 6.0 (-
7.0) um wide.
Isolated from basal portions of petioles and from
roots of Spathiphyllum sp. 'Clevelandii'’ in Apopka,
Florida. Deposited in the American Type Culture Col-
lection, 12301 Parklawn Drive, Rockville, Maryland
20002. Uso has ATCC, 44730.
DISCUSSION
Cylindrocladium spathiphylli is closely related to
CS floridanum but can be separated from the latter
because the conidia of C. spathiphylli are longer and
wider, the vesicle shape is globose (sphaeropedunculate
in C. floridanum), and lateral stipes are absent (pre-
sent in C. floridanum). Further, the teleomorphic state
of C. spathiphylli is wanting, whereas that of C.
floridanum is Calonectria kyotensis Terashita (1968).
LGesnould also be notéd that other isolates ‘of C-
spathiphylli obtained from various locations in Florida
were identical to the described type culture, which
clearly represents a distinct species.
ACKNOWLEDGMENT
The authors thank Dr. Walter E. Forehand, Chairman
and Professor, Department of Classics, the Florida State
University, Tallahassee, FL 32306, for rendering the
Latin diagnosis, Dr. James W. Kimbrough, Professor,
Department of Botany, University of Florida, Gaines-
wider rh. 32611 “and Dr. William’ J. Dress, ‘Professor
Emeritus, L. H.. Bailey Hortorium, Cornell University,
Ithaca, NY 14853, for their reviews, and Mrs. Janet
Temple for typing the manuscript.
REFERENCES
bell. D..K. and E. K. Sobers. 1966. A peg, pod, and
root necrosis of peanuts caused by a species of
Calonectria. Phytopathology 56:1361-1364.
Boedijn, K. B. and J. Reitsma. 1950. Notes.on the
genus Cylindrocladium. Reinwardtia 1:51-60.
Hansen, H. N. and R. E. Smith. 1932. The mechanism of
variation in imperfect fungi: Botrytis cinerea.
Phytopathology 22:953-964.
Zi2
Hansen, H. N. and W. C. Snyder. 1947. Gaseous sterili-
zation of biological materials for use as culture
media. Phytopathology 37:369-371.
Morrison, cR. Hie and (Dia W.-Erench. ~ 1969. slaxonomy, of
Cylindrocladium floridanum and C. scoparium.
Mycologia 61:957-966.
Schoulties, Ci. Land’ N.-Ek. Bl-GhoLi.) 19S). es Asroot
and foliar disease of Spathiphyllum 'Clevelandii'
incited by Cylindrocladium floridanum. Phyto-
pathology, 7 V2 255 (Abstr.)-
Schoulties, C. L. and N. E. El-Gholl. 1980a. Pathogeni-
city of Cylindrocladium floridanum on Spathiphyllum
sp..,.cv. Clevelandii<?.Procy, FlaweState otk. c0c.
93:183-186.
Schoulties, C. L. and N. E. El-Gholl. 1980b. Root and
petiole rot of Spathiphyllum sp. 'Clevelandii'
caused by Cylindrocladium floridanum. Fla. Dep.
Agric. Consum. Serv. Div. Plant Ind. Plant Pathol.
CLECs aNO nee [Dsus cp.
snell,s W.: H. and E.sA. Dick. 19/1. A.selossaryo&
mycology. Harvard University Press, Cambridge,
MA.
Sobers, E. K. 1972. Morphology and pathogenicity of
Calonectria floridana, Calonectria kyotensis, and
Calonectria uniseptata. Phytopathology 62: 485-
487.
Sobers, E. K. and C. P. Seymour. 1967. Cylindrocladium
floridanum sp. nov. associated with a decline of
peach trees in Florida. Phytopathology 57:389-
BOSs
Terashita, T. 1968. A new species of Calonectria and
its conidial state. Trans. Mycol. Soc. Japan
8:124-129.
MYCOTAXON
MOdeV Is NOs don DD. clomsoe October-December 1982
ERYNIA NEOPYRALIDARUM SP. NOV. AND CONIDIOBOLUS
APICULATUS » PATHOGENS OF PYRALID MOTHS
COMPONENTS OF THE MISDESCRIBED SPECIES,
ENTOMOPHTHORA PYRALIDARUM
[ZYGOMYCETES: ENTOMOPHTHORALES ]
ISRAEL BEN-ZE'EV*
Agriculture Canada, Research Branch, Research Station,
Harrow, Ontario NOR 1GO Canada
ABSTRACT
A reexamination of the species Entomophthora pyrali-
darum Petch showed that the original description was based
on different species, namely Conidiobolus apiculatus (Thax-
ter) Remaudiére & Keller and another species described here
aS Erynia neopyralidarum sp. nov. The name £. pyralidarum
Should, therefore, be considered as a junior synonym of c.
apiculatus.
Entomophthora apiculata (Thaxter) Gustafsson, the pre-
vious synonym of C. apiculatus, included Entomophthora
pseudococci (Speare) [now Conidiobolus pseudococci (Speare)
Tyrrell & MacLeod] as a synonym. Cc. pseudococci was sub-
sequently restored as an independent species because of pro-
duction of microconidia. The restoration of this species is
further supported in the present study by evidence of other
differences between c. apiculatus and C. pseudococci in CO-
nidial and rhizoidal morphology.
Entomophthora destruens Weiser & Batko, a pathogen of
mosquitoes, has rhizoids of the same type as C. pseudococci,
as well as other familial and generic conidioboloid char-
acters. A new combination, Conidiobolus destruens, iS pro-
posed for this species.
a
Visiting Fellow, National Science and Engineering Research
Council, Canada.
274
INTRODUCTION
Several species of Entomophthorales discovered by T.
Petch, including Entomophthora pyralidarum Petch, apparent-
ly were not encountered again, judging by their very infre-
quent subsequent citation in the literature. This fungus
was described (Petch, 1937) from dead "grass moths" (Lepi-
doptera: Pyralidae) found in December 1923 and January 1924
in three localities in Ceylon (Sri-Lanka). Specimens had
been sent for identification to Prof. R. Thaxter who stated
(fide Petch, 1937) that he was not acquainted with the fun-
gus.
The description of £. pyralidarum by Petch (1937) was
rather sketchy and was not accompanied by photographs or
drawings. Characters of taxonomic importance at the generic
level, such as the number of conidial nuclei and structure
of the conidial wall, were not mentioned. For these rea-
sons E. pyralidarum was not included in the classification
proposed by Batko (1964-1974, cited from Ben-Ze'ev and Ken-
neth 1982a) or in the neobatkoan classification of Humber
and Ben-Ze'ev (1981). Probably for the same reasons it was
not mentioned in the neobatkoan classifications of Remau-
diere and Hennebert (1980), and Remaudiére and Keller (1980).
Ben-Ze'ev and Kenneth (1982a) placed FE. pyralidarum in a
newly proposed group, Entomophthora nomina provisoria, a
group designed for species of Entomophthorales that are in-
completely described.
An examination and eventual complementary description
of E. pyralidarum appeared to be possible because most fun-
gi described and/or collected by Petch and Thaxter were
carefully preserved.
E. pyralidarum as depicted by Petch (1937) was an "anom-
alous" species. It possessed branched conidiophores and
rhizoids, features which are very frequently associated in
the genus Erynia Nowakowski emend. Humber and Ben-Ze'ev
(1981), and pyriform, oval or globose conidia which are
usually characteristic of other genera of Entomophthorales.
Globose primary conidia, however, have not been encountered
yet in any of the 47 species classified as Erynia by Nowa-
kowski (1881), Remaudiére and Hennebert (1980), Remaudiére
and Keller (1980), Humber and Ben-Ze'ev (1981) and Ben-
Ze'ev and Kenneth (1982b). This fungus, therefore, was
considered to be a good test of the view outlined by Humber
(1981), that "one of the most severe tests for a classifi-
cation system is to see how well it can handle the least
typical species from the group being classified."
215
MATERIALS AND METHODS
Specimens Examined
The following dried specimens were obtained from T.
Petch's collection, through the courtesy of The Herbarium,
Royal Botanical Gardens, Kew, England: package no. 9,
labelled "'Entomophthora pyralidarum Petch on moth, Kandy,
January 1924, per G. M. Henry, ex Herb. Petch"’, containing
a moth on a piece of leaf; package no. 10, labelled "'Ento-
mophthora pyralidarum Petch on moths, Vavuniya, Dec. 1923,
ex Herb. Petch", containing a moth and remains of a second
one; package no. 11, labelled "Entomophthora pyralidarum
Petch on moths (Pyralidae), Peradeniya, Dec. 25, 1923, ex
Herb. snaeretch", containing three moths and pieces of tree
bark to which they were attached by rhizoids.
Four slides labelled "Acc. no. 6423 Entomophthora api-
culata Thax. on Pyralid moth, Peradeniya, Ceylon, T. Petch
1924" were obtained from R. Thaxter's collection, through
the courtesy of The Farlow Herbarium, Harvard University,
Cambridge, Mass., U.S.A. The fungus in these slides was
identified by A. G. Kevorkian, about or previous to 1935
(Dr. D. M. Pfister, Curator, The Farlow Herbarium, pers.
commun.), which was prior to the description of £. pyrali-
darum by Petch (1937). The specimen from which these slides
were prepared was examined by Dr. R. A. Humber (pers. com-
mun.) and found to contain Conidiobolus apiculatus.
Microscopic Preparations and Measurements
Small portions were dissected from exsiccata and mount-
ed for microscopic examination on glass slides, in lacto-
phenol-cotton-blue (LPCB), in acetocarmine (Lee, 1950; Hall
and Bell, 1963) or in safranin-0 (Bandoni, 1979). Slides
were gently heated over an alcohol flame to enhance stain-
ing and to exclude air bubbles.
The ocular micrometer used for measurements had a cal-
culated error range of + 0.6 pm. Diameters of flattened
cylindrical structures were calculated using the formula
D = 2-width/T.
RESULTS AND DISCUSSION
The specimens collected by Petch in Kandy and Vavuniya
(packages nos. 9 and 10 from The Herbarium, Royal Botanical
Gardens) and the specimen collected by Petch in Peradeniya
and sent to Thaxter (slides obtained from The Farlow Her-
barium) contained the same fungus, Conidiobolus apiculatus
276
(Thaxter) Remaudiére & Keller. The specimens in Petch's
collection were still labelled as "Entomophthora pyrali-
darum'', indicating that they had not been reexamined,
whereas the slides in Thaxter's collection were correctly
labelled as Entomophthora apiculata (the synonym of Cc.
apiculatus used until 1980). Package no. 11 in the Petch
collection contained, side by side, one moth infected by
C. apiculatus and two others infected by a species of
Erynia. The moth infected by C. apiculatus was removed by
the present author from specimen box no. 11 and was placed
in a new box labelled "lla - Conidiobolus apiculatus on
moth, Peradenyia, Ceylon, Dec. 25, 1923".
The description of £. pyralidarum (Petch, 1937) con-
tains elements of C. apiculatus, together with elements of
the Erynia sp., and apparently some elements of one or more
Deuteromycetes which are also present in the specimens of
the Petch collection. Accordingly, Entomophthora pyrali-
darum Petch 1937 is considered as a junior synonym of Co-
nidiobolus apiculatus (Thaxter) Remaudiére § Keller (= Em-
pusa apiculata Thaxter 1888). The Erynia elements are des-
cribed here as a new species.
Erynia neopyralidarum sp. nov., NON Entomophthora
pyralidarum Petch 1937, Trans. Br. Mycol. Soc.
IA ote
Conidiophora primaria ramosa, digitata, determinata.
Conidia primaria uninucleata (? bitunicata), pyri-
formia, papillata vel papillata-apiculata (secundum
Lakoni [1919] systema), 16.0-28.0 x 12.0-17.2 pm,
papillae 1.2-4.6 pm altae; conidia secundaria co-
nidiis primariis similia sed minoria, minime 13.7
x 9.2 pm, in conidiophoris brevibus ex conidiis
primariis lateraliter orientibus portata. Sporae
perdurantes sphaericae, subhyalinae, diametro 25.2-
42.4 pm (med. 33.1 + 3.9 pm), sporarum pariete 2.3-
3.4 pm crasso, episporio leni; sporarum nuclei 2-7,
ellipsoideis 5.7 x 6.9-8.0 x 9.2 pm. Rhizoidea
filiformia, unihyphalia, aliquando ramosa, interdum
apicem versus incrassata; pseudocystidia non visa.
In Pyralidae (Lepidoptera) imagine, Peradeniya,
Ceylon, Dec. 1923. Typus: exsiccati no. 11 "Enxynia
neopyralidarum, Peradeniya, Ceylon, Dec. 1923", et
lamina per microscopium Exynia neopyralidarum de-
signata, Ben-Ze'ev, no. 11 (I - VIII), Reg. Bot.
Herb., Kew, Anglia.
Primary conidiophores are digitately branched, deter-
CaN
minate and uninucleate in the conidiogenous cells (Fig. 1).
Primary and secondary conidia are pyriform, uninucleate.
Their nuclei (stained with acetocarmine -- Fig. 2) are en-
tomophthoroid, relatively large (see dimensions and dis-
tribution in Table 1) spherical or more frequently ellip-
soidal. The results of this study are ambiguous with re-
gard to the conidial wall structure in FE. neopyralidarum:
a separable outer wall was observed in only three of 102
conidia screened (Fig. 3). Since E. neopyralidarum fits
all other generic features-criteria of the genus Erynia as
defined by Humber and Ben-Ze'ev (1981) and by Ben-Ze'ev and
Kenneth (1982b), its conidia would be expected to be bitun-
icate rather than unitunicate. A similar scarcity of bi-
tunicate conidia has been observed by the present author
(Ben-Ze'ev, in preparation) in other Erynia species col-
lected by Petch, although more bitunicate conidia were ob-
served in the other species. It seems very likely that the
unitunicate condition observed in the conidia of Erynia spp.
in the Petch collection is an artifact caused by the pre-
servation technique used by Petch, and that the bitunicate
condition observed in some of these conidia reflects their
true nature.
Dimensions of E. neopyralidarum conidia, and other co-
nidial and resting spore parameters calculated in the pre-
sent study and those given by Petch (1937) for £. pyrali-
darum are compared in Table 1. Conidia usually have acu-
minate papillae (Fig. 1) of apiculata-type [according to
Lakon's (1919) classification], with a conspicuous or, more
frequently, inconspicuous collar. On average, the papilla
occupies 15.5% of the total conidial length and approxi-
mately 1/3 to 1/2 of the total conidial width. The ranges
in length and width given by Petch for the pyriform or oval
conidia coincide with the upper part of the range found in
the present study (Table 1). Petch (1937) probably mea-
sured conidia on a moth collected soon after death and his
range probably represented primary conidia or primary co-
nidia with fewer secondary ones than were included in the
range found in the present study. The fungus in the moths
kept in Petch's collection, from which conidia were mea-
sured in this study, probably developed for a longer time,
producing more secondary conidia. The present findings
indicate that the length/width ratios of primary and sec-
ondary conidia of E. neopyralidarum are very similar making
distinction between the two classes of conidia very diffi-
cult. The maximum dimensions of secondary conidia overlap
the measurements of the primary ones and therefore, are not
278
distinguishable, whereas the minimum dimensions in the range
found, 13.7 x 9.2 pm, are the minimal dimensions of second-
ary conidia. These conidia are borne on short conidiophores
arising laterally from primary ones (Fig. 3).
Resting spores of Entomophthora pyralidarum were des-
cribed by Petch (1937) as ''...zygospores (7), spherical,
smooth, hyaline, 14-24 p diameter...''. Spherical struct-
ures matching this description were found in specimen no.
11 in the present study (Table 1) but they were probably
not entomophthoralean resting spores. They were very thin-
walled and stained uniformly in acetocarmine and safranin-
OQ without showing nuclei, vacuoles or lipid drops (Fig. 4a).
These spherical structures could belong to one of several
Deuteromycetes present in Petch's specimens as mycelium,
conidia, and possibly chlamydospores.
True entomophthoralean resting spores were found in
the present study in one of the two moths in package no. 11
from Peradenyia which contained the other structures of E.
neopyralidarum. They were found in mats of mycelium and
hyphal bodies but their mode of production (as zygospores
or azygospores) remains unclarified. These resting spores
were much larger than those described by Petch (Table 1)
and had a thicker wall consisting of a somewhat darker (sub
hyaline) episporium which was sometimes partly detached
from the lighter, apparently bi-layered endosporium (Fig. 5).
Resting spores stained in acetocarmine contained 2-7 nuclei/
spore. These nuclei were more frequently ellipsoidal than
were nuclei of conidia but were within the same range in
FIGS. 1-4: Enxynia neopynalidarum sp. n. on Pyralid moths,
photographs from the type material in T. Petch's
collection
1. Digitately branched conidiophores, x 500.
2. Primary (and possibly secondary) conidia stained with
acetocarmine, showing nuclei and variously shaped
papillae, x 1000.
3. Primary conidia with secondary conidiophores, two of
them showing secondary conidia; the conidial outer
wall layer is detached at the apex of the conidium in
the lower left corner (probably by the pushing conid-
iophore), x 1000.
4. Unidentified spherical structures (a) possibly those
interpreted by Petch (1937) as resting spores (arrows)
together with conidia (b) stained with acetocarmine,
x 500. Conidial nuclei are visible.
ames
280
TABLE 1. Quantitative features (parameters) of Exynia neopynralidarum.
Those measured or calculated in this study are from Petch's specimen no.
Lib
~~. | a. Pilik=@arg 0) “7 | Present: 2G) Marit: hia
Parameter n study n (1937)
Conidial length X
width (um)
min.-max. X 135 =217 aOmex 16-28 X
min.-max. RAS I/, 5) 12-16
x length (s; sz) X TOE VARIES! (AAS. OA) F. fy ?
xX width (s; sy) aM (IAG (OG IS),
Conidial length/
width ratio
min.-max. (xX) 102 1320-1-.880(L.os)
(Ss; Sz) (0.14-0.014)
Conidial papillar
height (ym)
min.-max, (most 102 1.15-4.60
frequent, 65% of (2.30-3.40)
measurements)
Resting spore dia-
meter (um)
min.-max. (%; s; s) L6" (25 8e—42 04 W335i
wall thickness (min.- 3.94; 0.96)
max.) 2.3-3.4
Spherical structures
considered resting spores
by Petch (1937) -— diameter
(um)
min.-max. (X; S; Sx) 37 8.0-22.9 (16.6; > 14.0-24.0
wall thickness Set P Oseyy) AsO a
Conidial nuclei (um) approx.
spherical min.-max. (x); % 6.9) 45%
ellipsoidal min.-max; %
82
Resting spore nuclei (um) approx.
spherical (diameter) min.- 69 20%
max.; % *
ellipsoidal (length xX Sie,
width) min.-max.; % 6.9-9. 80%
*
(2-7 nuclei/resting spore,
measured in 16 spores)
Size (Table 1). Hyphal bodies were usually short, unbranch-
ed, with few nuclei (Fig. 6). Chlamydospores, as described
by Petch (1937) have not been observed in the material con-
taining E. neopyralidarum. His description, however, seems
to fit some of the hyphal bodies of C. apiculatus during
formation of resting spores, as observed in the slides from
281
- 5-7: Exynia neopynalidarwm sp. n. on Pyralid moths, photographs
from the type material in T. Petch's collection
Resting spores stained with acetocarmine to show nuclei: (a) ina
mycelial mat, x 250; (b) x 500; (c) resting spore with 5 nuclei
and a partially detached episporium, x 1000.
Short, binucleate hyphal body (acetocarmine), x 1000.
(a) Monohyphal rhizoids showing ramifications (cotton blue), x 125;
(b) bifurcate extremity of a rhizoid with two funnel-like hold-
facts, x 500.
282
the Thaxter collection.
Rhizoids of E. neopyralidarum from specimen no. 11
were numerous, monohyphal, sometimes branched but not pro-
fusely so, with infundibuliform enlargements as holdfasts.
Their filaments were devoid of cytoplasm at maturity appear -
ing flattened in microscope preparations with a calculated
diameter range of 10.2-20.4 pm (Fig. 7). Pseudocystidia
were not observed by Petch or by the present author.
Type material of FE. neopyralidarum is preserved as
dried specimens and microscope slides at The Herbarium,
Royal Botanical Gardens, Kew, England. In addition, ori-
ginal negatives and photographs of temporary microscope
preparations are preserved at the same institution.
Conidia of E. pyralidarum described by Petch (1937) as
"...globose, 18-28 p diameter, with a conical papilla 4-8 p
high..."' (source 5 in Table 2) were not encountered in the
two moths of specimen packet no. 11 which contained EF. neo-
pyralidarum but were in a majority in the third one from
this packet. The nuclei of these globose conidia were not
stained by any of the stains used in the present study.
These conidia were similar in shape to those of C. apicula-
tus found in specimen packages nos. 9 and 10, although they
were Significantly smaller (source 7 in Table 2). Moreover,
the rhizoids on the moth containing these conidia were iden-
tical with those found in the other two packages (nos. 9
and 10) which contained C. apiculatus. These findings in-
dicate that the third moth in package no. 11 was infected
by C. apiculatus and that by the time it was collected most
of the primary conidia had produced secondary or tertiary
ones which are smaller than primary conidia.
A few conidia of FE. neopyralidarum, each with a clearly
visible nucleus, were observed in preparation no. lla, pre-
pared from the third moth in Petch's specimen no. 11, the
one infected by C. apiculatus. Because the three moths con-
taining the two different fungi were in the same package,
actually in contact with each other for many years, mixing
would be expected. This observation suggests how the two
different species became mixed in the description of E.
pyralidarum by Petch (1937).
E. neopyralidarum matches all the generic characters
of Erynia Nowakowski emend. Humber and Ben-Ze'ev (1981),
noting some ambiguity regarding the bitunicate condition of
its conidia. Because its subgeneric characters are incom-
pletely known, it cannot be included in any of the subgenera
of Erynia in the classification of Ben-Ze'ev and Kenneth
(1982b) and it is, therefore, placed in the temporary sub-
group Erynia sensu lato. By having pyriform conidia with
283
TABLE 2. Comparative conidial dimensions of Conidiobolus apiculatus,
C. major and C. pSeudococek: (1)--C. apiculatus combined from various
hosts, C. major on imago of Ptilodactyla serricokis (Coleoptera) -
from Thaxter (1888); (2), (3) and (4)--C. apiculatus from Diptera and
Psocoptera spp., (2) and (3)--C. major ona Tipuka sp. (Diptera) and
from a culture isolated from an aphid (Homoptera), respectively - from
Gustafsson (1965); (5)--C. apiculatus (as Entomophthora pynalidarum -
from Petch (1937); (6)--C. apicukatus on adult moth (Lepidoptera: Pyral-
idae) from Petch's specimen no. 9 of E. pynralidarum (prep. no. 9a, this
study); (7)--C. aptculatus, secondary or tertiary conidia on adult Py-
ralid moth from Petch's specimen no. lla, formerly E. pynrakidarum (prep.
no. lla this study); (8)--C. pseudococck on Pseudococeus calceolariae
(Homoptera), *() length and mean calculated here from Speare (1912, Pl.
Deets e Oe Cy at pik fy Ky!) s
Source C. apraculatus C. major
length x width (x) ym length x width (x) pm
(1) 30-37 x 28-30 (35 s¢ S10)» )) 55-60 x 38-45 ( ? )
(2) 25-39 x 21-30 (32 x 265 )) 40-64 x 37-55 (55 x 45)
(3) 2a=570 xe 2) (28 So CAS) SHAE) bie SiS ine CSO) Be AYA),
(4) 18-32 x 16-27 (26 xXEeZow)
(Gey) ? x 18-28 ( ? )
(6) 29.8-51.5x 20.6-45.8 (SO mex les)
(length s=3.14;
width s=3.21; n=72)
(7) HISAR She NOK ESS) (22,5 e Xess)
(length s=3.78;
width s=3.5; n=25)
C. pSeudococck
(8) * (25.5-34.4)x 20-25(32.3x23.9)*
(length s=2.5;
width s=1.9; n=7)
an average 1/w ratio of 1.5, which superficially resemble the
conidia of the Entomophaga grylly-type, Eneopyralidarum is
one of the least typical species in Erynia with regard to
conidial shape. However, the exclusion of the conidiobolo-
id elements changed the "'anomalous'' character of this spe-
cies, as it appeared to be from Petch's description.
Remarks on Conidiobolus apiculatus and Related Species
Empusa apiculata Thaxter was first described by Thaxter
(1888) in adults of various genera of Lepidoptera, in one
lepidopterous larva, in numerous genera of small flies and
gnats (Diptera) and in adult leafhoppers (Homoptera). One
of the isolates studied by Thaxter was from a beetle (Cole-
optera) and was similar to the other £.apiculata isolates,
except for substantially larger conidia. This isolate was
classified by Thaxter (1888) as E.apiculata var. major.
These two variants were transferred by Gustafsson (1965) to
Entomophthora as two independent species, E. apiculata
(Thaxter) Gustafsson and E. major (Thaxter) Gustafsson.
284
Recently the two species were transferred to Conidiobolus
by Remaudiére and Keller (1980) as C. apiculatus Rem. § Kell.
and C. major (Thax.) Rem. & Kell. Gustafsson (1965) repor-
ted isolation and growth of these species in artificial cul-
ture. He added Psocoptera (unidentified genera and spp.)
to the host-range of C. apiculatus, and gall-midges (Dipte-
ra: Cecidomidae) to that of C. major. “C. apiculacus, was
reported later to have killed aphids (Homoptera) in France
(Thoizon, 1970), and in Israel and South Africa (Ben-Ze'ev
et al., 1981, and unpublished data).
To summarize, Petch's (1937) description of Entomoph-
thora pyralidarum comprised: 1) branched conidiophores,
found in the present reexamination to belong to Erynia neo-
pyralidarum; 2) oval or pyriform conidia with conical pa-
pillae, demonstrated here to be uninucleate, probably bi-
tunicate and belonging to E. neopyralidarum; 3) globose
conidia with conical papillae, larger than the former ones,
demonstrated in this study to be Conidiobolus apiculatus;
4) resting spores considered here to be of unknown origin,
and demonstrated as different in size and cytology from the
genuine resting spores of E. neopyralidarum; and 5) rhizoids
which were insufficiently described by Petch (1937), and
which according to his sketchy description could be those
of either C. apiculatus or E. neopyralidarum.
According to the most recent interpretation of the In-
ternational Code of Botanical Nomenclature (13th Internat-
ional Botanical Congress, 1981) there were two possible re-
solutions to the classification of FE. pyralidarum: 1) to
exclude the conidioboloid elements and to propose a new com-
bination, with an emended description for the Erynia ele-
ments, under the specific name "pyralidarum"'; 2) to syn-
Oonymize E. pyralidarum with the senior specific name, C.
apiculatus, based on the conidioboloid elements, and to
FIGS. 8-10: Conidiocbokus apiculatus on Pyralid moths from
T. Petch's collection, specimens nos. 9, 10
and lla
8. Simple conidiophores extended toward their apices, the
two on the upper right corner with neck-like constric-
tions below the developing conidia, x 250.
9. Primary conidia (cotton-blue), the one on the extreme
right with an apiculate papilla germinating to produce
a secondary conidium, x 1000.
10. Characteristic monohyphal rhizoids, unbranched, with
irregularly discoid holdfasts, x 125, the one on the
extreme right, x 250.
260
286
describe the Erynia elements as a new species. The second
solution was chosen because: 1) most of Petch's specimens
contained C. apiculatus; 2) at the time that E. pyrali-
darum was described by Petch (1937) sufficient information
about C. apiculatus was available to easily avoid a misdes-
cription; and 3) the specimen sent by Petch to Thaxter was
already identified as C. apiculatus in 1935 and was avail-
able if Petch wanted to reexamine it before publishing his
findings from Ceylon in 1937.
Thaxter (1888) and Gustafsson (1965) described the co-
nidiophores of C. apiculatus, under previous synonyms, as
unbranched or slightly branched. The conidiophores of C.
apiculatus observed in this study were unbranched with char-
acteristic constrictions at the apices (Fig. 8) similar to
those shown by Gustafsson (1965, Fig. 33) for C. major.
The taxonomic importance of these constrictions was dis-
cussed by Humber (1981) and by Ben-Ze'ev and Kenneth (1982a) .
Primary, secondary and possibly tertiary conidia were mea-
sured from specimens nos. 9 and 1la and are compared in
Table 2. In both of these specimens the conidia had api-
culate papillae and the rhizoids were characteristic of C.
apiculatus (Figs. 9 and 10).
Gustafsson (1965) included Conidiobolus pseudococci
(Speare) Tyrrell §& MacLeod (under its former name, Entomo-
Phthora pseudococci Speare [1912]) as a synonym of £. api-
culata, because of similarity in conidial morphology, pre-
sence of rhizoids in both fungi and production of microco-
nidia by C. pseudococci and by one of Gustafsson's isolates
of "EZ. apiculata''. A comparison of conidial dimensions of
C. apiculatus, C. major and C. pseudococci.(Table 2) showed
that conidia of C. apiculatus and C. major from Thaxter's
study (1888) differed greatly in size. Conidia of C. api-
culatus in Petch's specimen no. 9 were slightly larger in
length and width than those of C. apiculatus described by
Thaxter (1888) and substantially larger than those of Gus-
tafsson's (1965) C. apiculatus isolates, but smaller by
approximately 25% than conidia of C. major. The size of
conidia did not allow a clear distinction between C. pseu-
dococci and Gustafsson's isolates of C. apiculatus. Remau-
diére et al. (1979) compared different isolates of Conidio-
bolus obscurus (Hall §& Dunn) Remaudiére § Keller and con-
cluded that there is substantial variability in the size of
conidia within and among species with spheroidal conidia
and that distinction according to this criterion is very
difficult among such species. Thus, the differences in co-
287
Nidialsize of the C. apiculatus isolates shown in Table 2
could be explained in consideration of the remarks of Re-
maudiére et al. (1979), or could be attributed to the pos-
sibility that Gustafsson's sources (3) and (4) were not
really C. apiculatus. This last possibility was discussed
by MacLeod and Miiller-Kogler (1973, p. 836) who suggested
that Gustafsson's (1965) isolates that had smaller conidia
than had Thaxter's isolate of FE. apiculata and that pro-
duced microconidia were probably related to either Conid-
iobolus pseudococci or to C. coronatus.
Thaxter (1888) and Gustafsson (1965) noted, without
providing measurements, that the ratio of papillar length
to conidial length of C. major was smaller than that of Cc.
apiculatus. Such conidial parameters as the absolute pap-
illar length (APL), proportional papillar length (PPL), co-
nidial length/width ratio (1/w), and apapillate conidial
1/w ratio (AC-1/w) were calculated in the present study and
evaluated as additional quantitative criteria to increase
precision of differentiation among species with spheroidal
conidia. A comparison of these parameters (Tables 2 and 3)
showed that C. pseudococci differs from both C. apiculatus
and C. major by having smaller conidia, a larger conidial
1/w ratio and a much larger PPL. C. major differs from C.
apiculatus by larger conidia (Table 2), larger resting
spores (Gustafsson, 1965) and a smaller PPL (Table 3: 14.34%
as compared to 16.10-17.38%). This apparently small differ-
ence in PPL between the two species is, however, substantial
enough to be readily detected in Thaxter's (1888) drawings
(Eineetowe ries. .05-70. Var. apiculata Vs. Figs. 7l@73,~var,
major)*. The APL allowed good distinction between C. major
and C. apiculatus or C. pseudococci but not between the
latter two species. The AC-1/w ratios for the three spe-
cies were about 1.0, meaning that their conidia without
papillae were almost perfectly spherical. Although this
ratio was of little value in differentiating these species,
it could be useful when species possessing spheroidal co-
nidia are compared with species that have ovoid or pyriform
conidia, e.g. C. apiculatus vs. E. neopyralidarum.
It is concluded that Gustafsson (1965) was justified
in elevating C. major to species level, but not in consid-
ering C. pseudococci to be a synonym of C. apiculatus.
*See Thaxter (1888, p.194-195, DESCRIPTIONS OF THE PLATES).
An error was made in the designation of drawings in his
Platenios
288
TABLE 3. Comparative conidial parameters of Conidiobolus apiculatus,
C. major and C. pseudococck calculated here from the following sources:
(1)--C. apiculatus from Thaxter (1888, Figs. 65-70), and C. major (l.c.
Figs. 71-73); (2)--C. apicukatus on adult Pyralid moth from Petch's
specimen no. 9 of Entomophthora pynakidarum (prep. no. 9a, this study);
(3)--C. pSeudococek on PSeudococcus calceolariae from Speare (1912, Pl.
Deel GSI poe y epee 7 ahi. TF) en
Parameter « APLC
(1) (2)
Absolute papillar length
(APL) min.-max. (x) um S09 we sa) 4.6-9.2 (6.4)
standard deviation; n LaGp OsOn UZ
Proportional papillar
length (PPL) as % of HY, SERIES 16.10%
average conidial length
Conidial length/width
Ta tEeuOm aly/we)
min.-max. (x) Piao (ie) Lol eGatlveo)
Se in Os09-N6 ORO6 7,
Apapillate conidial 1/w
ratio = length minus
papilla/width (AC-1/w)
min.-max. (X) 0.90-1.04 (0.99) 0.9-1.2 (1.04)
Syaer 0.05; 6 OO 2
C. mafonr C. psSeudococct
(1) (3)
APL: min.-max. (xX) ym 6.9=11 5 (9e 2) 623-80 (/53)
S70 Leon ON fae
PPL: % 14.34% 22 eLL%
l/w: min.-max. (xX) been a iow ad (Real 9) Lisa T. 4 itis)
SP 76 OAR Ss OaOae 7
AC-l/w: min.-max. (X) 0:95=0,.99" (0.97) 1 £OL—-1.09- (1.05)
Sron OOP Ts) 0.03757
In addition to the conidial differences, C.pseudococci dif-
fers from the other two species by ability to produce micro-
conidia and by different rhizoid endings (holdfasts).
Among the 31 species classified in the genus Conidiobolus
Brefeld (Ben-Ze'ev and Kenneth, 1982a) only C. apiculatus,
C. major, C. papillatus (Thaxter) Rem. §& Kell., and Cc.
pseudococci are known to produce rhizoids. These four spe-
cies are pathogenic to insects and their rhizoids are mono-
hyphal, but the holdfasts of the first three are similar
(Fig. 10), while those of C. pseudococci are different (Fig.
11b). Monohyphal rhizoids with holdfasts similar to those
of C. pseudococci are produced by some species of Erynia,
by Entomophthora culicis (A. Braun) Fres. (Gustafsson,1965;
Ben-Ze'ev, unpublished) and by Entomophthora destruens
289
FIG. 11: Conidioboloid rhizoids differing from those of
Contdiobolus apiculatus:
(a) distal extremities of Conidiobolus destuuens rhizoids,
redrawn from a drawing by Batko (in Weiser and Batko,
1966), and
(b) distal extremities of C. pSeudococct rhizoids redrawn
and brought to the same scale as (a) from a drawing
by Speare (1912).
Weiser and Batko (1966, Fig. 2) (reproduced here in Fig.
lla). The last species, which has conidioboloid nuclei
(Reva. Humber, pers. commun.) and all’ of the other generic
characters of Conidiobolus, was suspected for several
years to be synonymous with C. thromboides Drechsler (R.
Paani Der, sR: Oo. cOoper, Dewlyrrelljopers: <commun’. ja» How-
ever, the production of rhizoids and loricoconidia (conidia
which metamorphose into resting spores) described by Weiser
and Batko (1966) establish this species as different from
C. thromboides, whereas the conidioboloid characters justify
the proposal of the following new combination.
Conidiobolus destruens (Weiser §& Batko), comb.
nov. BASIONYM: Entomophthora destruens Weiser
and Batko, Folia Parasitol. (Praha) 13: 145-146,
Text -t1oye t—-ge tis. a! (Fig. 1-3) 62 (Pigwia2).
1966.
Since C. destruens does not produce capilliconidia or
microconidia, producing only Type I secondary conidia, it
290
is placed in the subgenus Conidiobolus subg. Conidiobolus
as defined by Ben-Ze'ev and Kenneth (1982a), close to the
other three rhizoid producing species classified there, Cc.
apiculatus, C. major and C. papillatus. Phylogentically,
however, C. destruens appears to be closer to Conidiobolus
(subgen. Delacroixia) pseudococci, because it has similar
rhizoids. Other species placed in subgen. Conidiobolus,
because they produce only Type I secondary conidia, appear
to be related biochemically to either subg. Delacroixia
(Sacc. §& Syd.) Tyrrell §& MacLeod or to subg. Capillidium
Ben-Ze'ev and Kenneth (Tyrrell and MacLeod, 1972; King,
1976; Ben-Ze'ev & Kenneth, 1982a, p. 422 and 428-431).
These phylogenetic problems encountered with species of the
subgenus Conidiobolus suggest that this subgenus is phylo-
genetically artificial, undoubtedly containing some species
that are more closely related to the other two subgenera
but that have lost the ability to produce microconidia or
capilliconidia. From the systematic point of view it is
convenient to retain the subgenus Conidiobolus until addi-
tional knowledge allows broader definitions for the other
subgenera of the genus Conidiobolus.
ACKNOWLEDGEMENT
I am thankful to the Royal Botanical Herbarium, Kew,
England, and to the Farlow Herbarium of the Harvard Univer-
sity, Cambridge, Mass. U.S.A. for loans of material from
the collections of T. Petch and R. Thaxter, respectively,
and to D. H. Pfister, Curator of the Farlow Herbarium for
valuable information about the specimens sent by Petch to
Thaxter.
I wish to thank B. E. Tucker of Washington University,
Seattle, Washington, U.S.A., R. P. Jaques and W. R. Jarvis
of Agriculture Canada, Research Station, Harrow, Ontario
and R. G. Kenneth of the Hebrew University of Jerusalem,
Israel for their constructive reviews of the manuscript,
and D. Stuntz of Washington University, Seattle, for re-
viewing and correcting the Latin description.
I am thankful to R. A. Humber and R. S. Soper, USDA,
Boyce Thompson Institute, and R. P. Korf, Cornell Univer-
sity, Ithaca, New York, U.S.A. and D. Tyrrell, Environment
Canada, Forest Pest Management Institute, Sault Ste. Marie,
Ontario, Canada for valuable correspondence and personal
communications.
vgs
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Mycologia 71: 873-874.
Batko, A. (1964). On the new genera: Zoophthora gen. nov., Triplo-
sporium (Thaxter) gen. nov. and Entomophaga gen. nov. (Phycomy-
cetes: Entomophthoraceae). Bull. Acad. Polon. Sci., Ser. Sci.
Biol iat S2g7520.
Ben-Ze'ev, I. and Kenneth, R. G. (1982a). Features-criteria of taxon-
omic value in the Entomophthorales. I. A revision of the Batkoan
classification. Mycotaxon 14: 393-455.
Ben-Ze'ev, I. and Kenneth, R. G. (1982b). Features-criteria of taxon-
omic value in the Entomophthorales. II. A revision of the genus
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Ben-Ze'ev, I., Kenneth, R. G. and Bitton, S. (1981). The Entomophthor-
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Hall, I. M. and Bell, J. V. (1963). Aceto-carmine — a rapid stain for
Entomophthora species. J. Insect Pathol. 5: 265-266.
Humber, R. A. (1981). An alternative view of certain taxonomic criteria
used in the Entomophthorales (Zygomycetes). Mycotaxon 13: 191-240.
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King, D. S. (1976). Systematics of Conidiobolus (Entomophthorales)
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and H. W. Beams), 1l-th edition, The Blakiston Company, Philadel-
phia.
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tomophthora species with pear-shaped to almost spherical conidia
(Entomophthorales: Entomophthoraceae). Mycologia 65: 823-893.
Nowakowski, L. (1881). O grupie owadomorkow (Empusaceae). Dzienn. III
zjazau Lek. Przyr. Polak. Krakow, Sekc. Bot. 6: 6/7.
Petch, T. (1937). Notes on entomogenous fungi. Trans. Br. Mycol. Soc.
21: 34-67.
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Entomophthora aphidis Hoffm. in Fres. Description de deux nouveaux
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Boston Soc. Nat. Hist. 4: 133-201.
ZO
Thoizon, G. (1970). Spécificité du parasitisme des Aphides par les
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Entomophthora destruens sp. nov. (Phycomycetes, Entomophthoraceae).
Folia Parasitol. (Praha) 13: 144-149.
MYCOTAXON
WOOP Te Now li opps (20552 5 October-December 1982
_ ire rt
SURVEY OF THE ARGENTINE SPECIES OF THE GAWODERMA
LUCTDUM “COMPLEX.
Mariaese: Bazzalo- & Jorge E. Wright¢
Departamento de Ciencias Biologicas,
Facultad de Ciencias Exactas y Naturales,
Universidad de Buenos Aires.
SUMMARY
Macro and micromorphological and cultural studies showed that the
Ganoderma Luetdum-complex is represented in Argentina by Ganoderma
luetdum s. str., G. oerstedit, G. resinaceum, G. zonatum and G. sub-
ambotnense var. lLaevtsporum var. nov. based mostly on the study of
type materials. A general key is presented in this complex and add-
itional synonyms for G. restnaceum are proposed. G. tuberculatum is
a synonym of G. oerstedtt.
The species of the Ganoderma lucidum (Leys.:Fr.)Karst.
complex are important wood-rotting fungi and cover a wide
array of hosts in Argentina: Tipuana, Acacta, Robinta, Oco-
ved, Pinus, Quercue, Ulmus, Acer, etc., inflicting serious
damage both to native and introduced trees. The group is
formed by those species possessing a typical "hymenodermis"
(Steyaert; 1972).
Unfortunately they have been poorly studied, particular-
ly for Central and South America. Thus, we lack modern keys
HOtecnetrandentitioat ion wcires Murrell), 1915s Dennis, 1976.).,
and those that are available are not sufficiently comprehen-
Sive and are based almost entirely on macromorphological
features. A study was thus carried out on most of the spec-
ies of the area having a hymenodermis, particularly on Ar-
gentine forms. This has allowed us to develop a tentative
key of the representative species and varieties from Central
and South America, with major emphasis placed on the basid-
iospores
The genus Ganoderma was created by Karsten (1881) based
ON IFotyporus lucizdue Leys .:Fr., the type of which is net
extant. It comprises those species of polypores whose pileus
and stem (when present) are covered by a crust with the
consistency and appearance of lacquer, sometimes brilliant,
Formerly Fellow of the Instituto Forestal Nacional, Argentina.
2 Member of the "carrera del investigador cientifico", Consejo Nacional
de Investigaciones Cientificas y Técnicas, Argentina. This paper
is part of a project on xylophilous Basidiomycetes of Argentina,
sponsored by the Consejo.
294
and with ovoid to ellipsoid basidiospores varying from
yellow to brown, provided with a hyaline perisporium that
is usually smooth and truncate, and a thick endosporium
with projections reaching to the external layer (Heim, 1962;
Furtado, 1962, 1965). The apex becomes truncate upon matur-
ation due to the collapse of the apical papilla. Until re-
cently the genus was included in the Polyporaceae but Donk
(1933) segregated the subfamily Ganodermoideae which he later
gave family rank as Ganodermataceae (Donk,1964) .
Since Patouillard (1889) revised the species to that
date, several workers dealt with the genus, describing
numerous species, among them Murrill (1908b, 1915), Lloyd
(OIA 1915 191731920, 91921, 1924)).) Torrend G19 coun em
gazzini (1926), Imazeki (1939), Rick (1938), and mainly
Steyaert (196la and b, 1962, 1967b, 1972). Steyaert's un-
timely death prevented him from writing a monograph of the
genus (but cfr. Steyaert, 1980).
In spite of the work carried out by these workers, the
taxonomy of the genus has been, and still is, rather chaot-
ic, principally. due to the great variety of forms encount-
ered and the different criteria used for classification.
It is now obvious that macromorphology alone is useless as
a basis for distinguishing among species.
New criteria arose after the studies of . Humphrey & Leus
(1931) and of Haddow (1931). The former though referring
only to the G. applanatum-complex (another difficult prob-
lem), emphasized the importance of employing anatomical
characteristics in their classification. Haddow (1931) was
the first to use the structure of the fruit-body's dermis
for the identification of a few species, but did not prop-
ose any descriptive terminology; he did, however, refer to
the spore ornamentation as the “smooth type" in Gunoderma
sesstle (=G. restnaceum) and the “rugose type" in G. luetdum.
This was later confirmed by Steyaert, (1962, 1972). .Fomour
species concept we have emphasized the type of spore ornam-
entation, shape and size, the number of pores per mm and,
in some instances only, the colour of the context. Thelatter:
has been widely used by previous authors but is not always
a reliable feature. Studies with the SEM corroborated the
structure of the spore wall proposed by Furtado (1962) as
well as the types of ornamentation proposed by Steyaert
(1972), to which we have added the "semirugose type" inter-
mediate between the typically “smooth" and"rugose" types.
Steyaert (1962, 1972, 1980) considered that.the different
forms constitute constellations or "complexes" according
to the types of dermis, which were defined and carefully
studied by Furtado (1965). All the species here described
fall in Steyaert's (1980) subgenus Ganoderma, Section Gano-
derma, although some related ones included in the key do not,
MATERIALS AND METHODS
The description of the basidiomes was made according to
their macroscopic features (size, colour, number of pores
per mm, length of tubes, colour and consistency of context,
presence or absence of stem, etc.). The microscopic feat-
ures studied were the types of hyphae, and the vegetative
and reproductive structures, mostly using the indications
given by Teixeira (1956). The hyphal system was studied by
dissecting small portions of the context and/or the dissep-
iments. Dermic elements were examined and measured in thin
sections perpendicular to,the pileus surface. Spore ornam-
entation and gasterospore features were studied both with
the optic and SE microscopes.
Cultural studies "in vitro" were made of some species ac-
cording to the methods of Nobles (1948, 1958, 1965), and
code numbers obtained.
Colours are according to Maerz & Paul (1930), and herbar-
ium abbreviations those of Holmgren & Keuken (1974).
KEY TO REPRESENTATIVE SPECIES OF THE GAWODERMA LUCIDUM-COMPLEX TO
BE FOUND IN CENTRAL AND SOUTH AMERICA.
tee SpOcenOrnamenta tion. GiStINCELY. YUGOSE®.. s.cl.scis o's eleien Galvre« escie e's 2
Ge OO ReROUNeMel cat 101 MOUs SO ss c15 c\' afer a «dose ole eve ole dime pie dseees wih 3
2. Context dark brown; spores 9-13 x 5-6,9 um..... G. tuetdum s. str.
2'.Context almost white; spores broadly ellipsoid,
Be UE ANN ls fk Sige Ie 21m ean omg ate eo G. punettsporum Furtado
3. Spores of the "smooth" type (with numerous slender
endosporic pillars that do not show up on the pe-
risporium, thus appearing smooth when observed with
PpemOmc te aR VCLOSCODe 0 LiV400 X) saws s.c ioe ci bale nw cacaineees « 4
3'.Spore ornamentation "semirugose, intermediate bet-
GC MME ON MAING S ai ees ia « teckel cee ore Vans oe (hago Ae eee oath ao 8
4. Gasterospores present in context and dissepiments.......... 5
MS OPO SMOMES MADSEN Cs dare et terol: ayes Wi otc eich o tails kee 3 Ga e Baran 6
9. Gasterospores with walls formed by veins or ribs
(particularly evident with the SEM); Brazil...
era a G. subambotnense P.Henn.
var. subanbotnense
9'.Gasterospores with smooth walls; Brazil and NE
(RST BTS Ns ak 0 aR on ee a oe a G. subambotnense P.Henn.
var. laevtsporum var. nov.
6. Pores 3-5 per mm; spores ellipsoid, 9-13 x
SEE TTT RRA IN il a earns G.restnaceum (Boud.)Pat.
De rOmesaustial ly parger than. 5 per MMs aa ales-cysss.ce a oacea ee tw on 7
7. Pores 5-6 per mm; spores 7-10 x 5-7 um, context
MO SMW UL Cetera tye as iN Lied bee alia hares G. parvulum Murr.
7'.Pores 6-7,5 per mm, mostly 7 per mm, spores
P=10-x 5=/ um, ‘context brown,....... .% G. btbadtostratum Steyaert
1 ;
Term used in the sense of Steyaert and employed much earlier by
Spegazzini for imperfect spores on or in normal basidiomes.
296
Bo CONTE X Cerda ake DIZOW Mladic: oi: sce ice cone fe farmer cul once meen te eerie 9
GY ACONLeXt almOStGIWiLterol, at Leds O.. nae
dark brown; basidiomes generally small
and sessile; spores 9-11 x 6-8; Mexico
aaces Cusastiees G. sesstitforme Murr.
9. Spores broadly ellipsoid, 9-14 x 6-10 um;
pores 4-7 per mm, surface of pileus usual-
Eyes tgonahy cuberCcuUlale..s «nce. G. oerstedit (Fr) Torr!
9'.Spores narrower, pileus surface not distinct-
Tye CUD GY CU: 8 Cero e src ues wie 3nsp.e ners oh ce eke ie: si imeenaen cae amen 10
10. Spores 11-13 x 5-7 um; pores 4-5 per mm;
Surface. of pileus smooth or concéentric-
Overs Ca Ce. aheteet iy wentiowa ) sesuean. Ge Zona cumenurn.
10'.Spores 10-11 x 5-6 um; pores 5-6 per mm;
surface of pileus radially rugose and 1
CONncentrical py sul Cates 7.0. G. subforntcatum Murr.
GAL NSO) DER MA Pikarst., Rev Mycol. 3ie( 9) eee oe
emend Patouitliard, Bul. Soc. Mycol. France 5: O77e cog
Xylophagous polypores producing a white-rot in wood. Bas-
jidiome annual or biannual, rarely perennial, pileate. Pileus
with an eccentric stem, or sessile and then dimidiate, rare-
ly centrally stipitate. Pileus and stem surfaces (when pres-
ent) covered with a crust with the consistency and appear-
ance of lacquer which may or not be shiny like varnish, well
defined, generally brilliantly coloured. Context corky, fib-
rous-coriaceous, or somewhat woody, more or less tomentose
or fibrillose, sometimes very hard and durable, with various
hues of brown, frequently whitish, sometimes with a thin
dark brown layer among the tubes. Hymenophore tubular, in
one or several layers, generally dark, with a smooth surface.
Pores small, mostly circular, ‘almost white, exceptionally
golden yellow. Hyphal system trimitic with hyaline, septate,
clamped, thin-walled generatives, usually branched and more
frequent at the margin. Skeletal hyphae aseptate, thick-
walled, light brown, long, wavy, scantily branched,occasion-
ally with dendritic endings, generally more frequent in the
context. Binding hyphae scarce, strongly branched and tort-
uos , aseptate, thick-walled, more slender than the skelet-
als. Cystidia and setae absent. Basidia short clavate to
napiform, 4-spored. Spores ovate-ellipsoid, with a truncate
apex and a t wo-layered cyanophilous wall, the external
smooth and hyaline, the internal brown, verrucose or retic-
ulate.
Type species: Ganoderma Luctdum (Leys.:Fr.)P.Karst.
Both yee belong to Section Characoderma, according to Steyaert
(1980).
297
GANODERMA LUCIDUM (Leys.:Fr.)P. Karst., Rev. Mycol. 3 (9): 17.1881.
= Boletus luctdus Leys. Fl.Halensis p. 300. 1783. = B. dimidia-
tus Thunb. , Fl. p. 348, tab. 39. 1784 (Fide Imazeki). = 8. laced
tus Timm, Fl. megalop. Prodr., p. 269. 1788 (Fide Imazeki). cs
sVrolLyporus Lucrdus LCYS <i hus s¥St. Mycol. [2 353. 1821: -=°P.
laceatus Timm:Pers., Mycol. europ. 2: 54. 1825 (Fide Imazeki).
= P. japonicus Fr., Epicr. p. 442. 1838 (Fide Imazeki)! #27. Zu-
etdus (Leys.:Fr.)Sacc. ,Syll. Fung. 6: 157. 1881. = Placodes lu-
etdus (Curt.:Fr.)Quél., Fl. Mycol. p. 399. 1888 (Fide Domanfski).
=F. japonicus FY. in sacc., Syll. hung. 6% 156: 1888 (Fide
Imazeki). = Ganoderma pseudoboletus(Jacq.) Murr., Bull. Torrey
bot. Cl. 29: 602. 1902 (Fide Imazeki). = G. laccatum Pat. in
Bresadola, Icon. Mycol. 21, t. 1004. 1932, non G. laccatum Bourd.
& Galz. 1928. Icon. Michael & Henning, Pilzfr. 2, t. 73. 1960
(Fide Domanski et al.).
Figs: 1-4, 23-25, 61.
Annual, sessile and dimidiate or, more frequently with
a lateral stem and then usually reniform (Figs. 1,3). Pilei
tsohated, small tao medium sizéd, 2-8 x 2-4,5 x 0.5-2 cm.
Pileus surface radially rugose and concentrically sulcate,
Deairantly slaccate, light reddish brown (P12 (@ Lsl2 of
Maerz & Paul), to dark reddish brown or mahogany (Pl. 7 L
6). Margin sterile, generally thick and blunt, sometimes
acute, white in actively growing specimens, becoming yellow-
ish and reddish brown inwards; in older specimens of the
same colour as pileus surface, and then incurved. Stem lat-
Srdievertical,acylindric, usually tong, slender. tortuous.
4=10 cm long;,0,5-2 <cm thick, reddish black to almost black,
laccate, brilliant, somewhat thicker at the base. Cutis
than, brilliant. black. Context. almost as thick as the tube
layer but thickening towards the base of the stem, ochrac-
eous brown when young (P1. 13 H 10) to dark brown when mat-
Uremirl 14 LM) corky. Dermis of thes"hymenodermis”™ type,
composed of thick, golden walled, claviform elements orig-
inating from the ends of skeletal hyphae, with narrow lumi-
na and blunt ends, arranged in a palisade-like hymenium
GFign 62)... 5-10 um thick. the total thickness of the dermis
14-42 um. These elements are covered by a thick layer of
a lacque-like substance that dissolves in a hot solution
of 5% KOH. Hymenophore white to yellowish white when young,
greyish white in mature specimens (Figs. 2, 4) with a tube
Lavyereupeco 7 mn long, slightly dighter than, context sGP 1%
13 D 8). Pores small, round, somewhat irregular, 4-7 per
mm, 6-200 um diam. Dissepiments 17-116 am diam. Hymenium
not persistent, composed of scant globose to subglobose
basidia, 9-19 x 7-14 um. No other hymenial elements pres-
ent. Basidiospores subovoid with the apex truncate, peris-
porium hyaline, smooth and thin, and endosporium golden
with relatively scant endosporic pillars, wide and long,
reaching the perisporium and rumpling it so that it appears
Scrongry .rugose (Figs..23=25. 61); 9-13 “x 5-6,9 um-> Hyphal
System trimitic with hyaline,thin-walled,clamped,septate
generatives, 1-4um diam. ,septa restricted to clamps, scant-
ilybranched, abundant at the growth margin of pileus and
dissepiments, rare or absent in the context (Fig. 71). Skel-
etals "arboriform" (Teixeira, 1956), aseptate, clampless,
very long, 3-6 um diam., scantily branched, branches with
limited growth at distal end, with thick golden walls, some-
times subsolid; they compose most of the context and dissep-
iments, originating immediately behind the growth margin
from generative hyphae (Fig. 68). Binding hyphae of the "Bo-
vista" type (Cunningham,1946a), aseptate, clampless, profuse
ly branched, tortuous, of limited growth, generally thinner
and lighter than the skeletals, 1-3 um diam., rather scant
and only present in the context; they are intertwined with
aa latter, giving the context its firm cohesion (Figs. 69-
LOR:
Hosts: at the base of trunks and on roots of harwoods,
rarely on conifers. Recorded also on Acer, Quercus, Casta-
nea, Alnus, Populus,\Fagus, Fraxtnusand Pinus (Domanski,
1967). According to Domafski (loc. cit.) it grows saprophy-
tically and only exceptionally attacks living trees -.boyee
(1938) agrees with this. However, Pirone (1957) through ex-
periments with Acer proved that it is an important parasite
of hardwoods.
Distribution: apparently worldwide in temperate and tropic-
al zones.It has been recorded for Europe, Asia, Philippines,
Australia, Africa and North and South America (in the latter
for Venezuela, Brazil, Uruguay and Argentina).
Apart from the records given below for Argentina, Spega-
zzini cited collections from the provinces of Cérdoba and
Chaco. We have been unable to find materials from those
areas.
Material studied: ARGENTINA: Buenos Aires: Don Torcuato, leg.
Kohn, 10.1X.1970 (BAFC 24406); Pereyra Iraola, IV.1971 (BAFC 24421);
Bella Vista, leg. Gallardo, 31.1.1965 (BAFC 25531). Capital Federal:
Jardin Botdnico, leg. Molina, 7.1V.1978 (BAFC 24408). Corrientes: 4 km
E of Paso de la Patria, leg. Krapovickas, 20.V.1966 (BAFC 24428). Mi-
siones:Parque Nacional Iguazu, leg. Bazzalo, 14.1I1.1980 (BAFC 25055);
Garupa, leg. Cricel, 16.1V.1976 (BAFC 24433); Arroyo Yacuy, leg. Gomez,
I 1.1969 (BAFC 24418). Tucuman: road to Tafi del Valle, km 19, leg.
Gémez, 22.V.1966 (BAFC 24419); road to El Cadillal, Hway 9, leg. Ruiz,
26.1V.1973 (BAFC 24425); ibid., leg. Guerrero & Bettuci, 27.1.1965
(BAFC 25524); no locality, leg. Spegazzini, VI.1917 (LPS 24886).
BRAZIL: Sao Leopoldo, leg. Rick (sub.G.rentdens) (BAFC 25599). FRANCE:
Basses Pyrénees, leg. Candousseau, 1972 (BAFC 25600). URUGUAY: no loc-
ality, leg. Berg (LPS 24851); Montevideo, leg. Arechavaleta, IX.1898
(LPS 24972); ibid., Cerro Largo, leg. Felippone (LPS 24882).
Figs. 1-4: Ganoderma lucidum s. str. 1-2: aspect of bas-=
idiome; 3: surface of pileus; 4: hymenophore. Figs /) 3-0.
Ganoderma subambotnense var. Llaevtsporum. 5: surface of
pileus; 6: hymenophore (all figures 1/2 x).
300
Cultural features Figs. 75-79, 81-82, 88-90, 107.
Strains: BAFC n°112 = ARGENTINA: Catamarca, Dique de Collagasta
leg. Laterra, 14.1.1981. BAFC n°815= ibid.
Code Numberii2.632°S. 10 47375 9) 40.53..704e on
Macroscopic characters: growth slow, covering Petri dishes in 5
weeks. Mycelium mat not dense, adhering to agar, arranged in concentric
bands the lst week, alternatively white and ochraceous (Pl. 11 K 1).
After the 4th week it becomes totally ochraceous yellowish with more or
less darker concentric bands (Fig. 107). Texture completely farinaceous,
margin subfelty. Reverse:discoloration brown. Margen regular, border
smooth. Odour sweetish. Oxidase reaction: with tannic acid= ++++ with
growht; with gallic acid= ++++ without growth; with gum guaic = +.
Microscopic features: marginal mycelium formed only by generative,
clamped hyphae, with septa restricted to clamps, scarcely branched,
2,5 um diam. (Figs. 88-90); during the 2nd week fibrous hyphae appear,
Slender, hyaline, thick-walled, clampless, heavily branched, 1-2 um
diam. (Figs. 77-78). Ochraceous yellowish mycelium formed by: i) gener-
ative hyhae as above; ii) clamped, thick-walled hyphae, golden, un-
branched, 2-7 um diam., of uniform thickness or like rosary beads
(Figs. 79, 81-82); iii) stag-horn clamped hyphae, thick-walled , with
numerous branches, generally dichotomic, projecting in several planes,
1-3 wm diam., or with short branches laterally disposed in a single
plane at regular intervals, 2-3 um diam.; iv) globose, thin-walled
golden yellow cuticu lar cells which are very abundant, tightly packed
forming a “pseudoparenchyma", 5-20 um (Figs. 75-76); v) fiber hyphae
as in the margin, scarce. Submerged mycelium similar to aerial.
These cultures are characterized by a strong yellowish
colour that extends to all the mycelial layer, whereas in
other species it is limited to a few spots and is not so
intense. The cultures resemble those of G. subambotnense
var. laevisporum but differ in the lack of chlamydospores.
For a long time the G. Zuetdum-complex has been a diffic-
ult problem for taxonomists, It has been a traditionmtuoecon=
sider all the stipitate forms as ¢.° luetdum’s. etry aneuene
sessile ones as G. restnaceum (= G. sesstle Murr.). This
distinction has an absolute lack of anatomical foundation.
When one deals with a krge number of specimens, clear dis-
tinctions appear with regard to spore ornamentation, what-
ever the macroscopic configuration may be. This had already
been observed by Haddow (1931) and was only reconfirmed 35
years later by Steyaert (1967a). Haddow termed the spores
Of G. Luctdum as of the "rugose" type. Unfortunately
the holotype of this species has not been found, although
Steyaert (1972) states there is a coloured illustration of
such specimen in Flora Londinensis (1781), which was collec-
ted at Peckham. S of London; This would constitute thespnes=
Figs. 7-8. Ganoderma sessile (Holotype). 7: hymenophore;
8: surface of pileus, 9-10. G. resinacewn, BAFC 24450. 9 hymenophore;
10: surface of pileus. 11-12. G. subambotnense var. laevtsporum.
11: aspect of the basidiome; 12: hymenophore (all figures 1/2 Xs
Ta
Sz
ent type of the species. However, such an illustration does
not reveal the spore features. Attempts to find a neotype
at Peckham have failed. Karsten, the founder of the genus,
left at H a specimen with the same type of "rugose" spores
(Steyaert, 1972), which could be selected as neotype and
thus arrive at a satisfactory "modus vivendi" to distinguish
among both species.
GANODERMA SUBAMBOINENSE P. Henn. Fungi amazonici I. Hedwigia
43: 175. 1904 (sub Fomes (Ganoderma) subamboinensts) var. LAEVISPORUM
NAS fe Ge = MANOA RA
Figs. 5-6, 11-12, 46-50, 62-71.
A typo dtffert gasterosports laevts. Holotypus Argentina, Buenos
Atres, Tigre, leg. Connon, 15.V.1980, tn Herb. BAFC 25525 conservatus
est.
Annual, sessile, dimidiate or, more frequently, stipitate
flabelliform to conchate, pleuropus, with isolated pilei
which are sometimes concrescent, small to medium-sized, 3,5-
13 x 2,5-10,5 x. 0,2-1,5..cm.. Pileus surface radially enueoce
with concentric, slightly deep sulcations, or radially rug-
ose and then with numerous, deep, concentric sulcations,
that make it appear undulate (Figs. 5, 11); dark reddish
brown (Pl. 7 L 11 of Maerz & Paul), gradually becoming light
er towards the margin, with a narrow yelbwish -orange band
(P1. 1130 7) in young specimens; in mature ones this grad-
uation is not observed, the colour tending to become homog-
eneous, brilliantly laccate. Margin sterile, straight, yel-
lowish white (PT. 11) Bul), thin and acute’ to thie cand
blunt; in the first case undulate and somewhat irregular;
in the second smooth and regular. Stem horizontal, short and
thick or long and slender, tortuous, reddish black to almost
black, “Laccate,; brilliant, cylindric. 15-5. 5 "cn pond. aoe
2,5 cm thick. Section very thin tn general, sligqneglyoswourm
en at the base, 0,3-1,5 cm deep at about half the radius.
Cutis thin, blacks brilliant. Context stn) hepa osrare lm
thick, light brown, almostewhite {Pls iy e Seca,
darker in a narrow zone above the tubes; in the Brazilian
specimensbrown (Pl. 13 H 8); corky. Dermis of the "hymeno-
dermis" type, composed of clavate elements with thick walts
and blunt ends, 5-10 um diam., covered by a thick layer of
lacquer that dissolves in a hot solution of KOH (FiqumGc.
16-36 um thick. Hymenophore poroid, concolorous with margin,
tube layer 1-4 mm long, concolorous (Figs. 6, 12); pores
circular, 4-7 per mm, 98-260 um diam., dissepiments 36-170
um. Hymenium formed only by easily collapsing subglobose ba-
sidia, 4-spored, 9-13 x 7-14 um (Fig. 57). Basidiospores
broadly ellipsoid with truncate apex (Figs. 46-50), of the
"smooth" type, perisporium hyaline, thin, endosporium light
Figs. 13-16. G. oerstedit. 13-14: aspect’ of the basadie-
me (BAFC 24441); 15-16: aspect of the basidiome (Holotype of
G. tuberculosum ) (both 1/4 x). Figs. 17-22. G. zonatum. 1/-
19: aspect of the basidiome of the, Holotype (1/2 x); 20-22%
aspect of the basidiome of BAFC 24416 (1/2 x).
304
brown, thick, endosporic pillars numerous, slender, not af-
fectingthe perisporium; small, 6-9 x 4-6 um. Hyphal system
trimitic with clamped, thin-walled, hyaline generatives,
with septa restricted to clamps, sparingly branched, 1-3 um
diam., abundant at the growth margin of the pileus, but also
present in the context and dissepiments (Fig. 71). Skeletals
aseptate,:' thick-walled, almost hyaline, solid to subsolid,
arboriform at the end of branches, 3-8 um diam., they form
the bulk of the context and dissepiments (Fig. 68). Binding
hyphae of the "Bovista" type, thinner than the skeletals,
almost hyaline, thick-walled, aseptate, heavily branched,
1-3 am diam., only present in the context (Figs. 69-70).
Gasterospores abundant in the context, ‘present in’ tne dissep
iments originating from vegetative hyphae, with dense cont-
ents and thick-walled, sometimes with 1-2 guttulae, almost
spherical, completely smooth, 7-14 x 6-12 am (Figs. 63-6778
Hosts: on dead fallen trunks of various Species a onmeuic
Platanus (at considerable height on the trunk), and on
stumps of Prnus taeda.
Distribution: Brazil, Argentina (Buenos Aires, Misiones).
Material studied: ARGENTINA: Buenos Aires: Tigre, leg. Connon, 15.
V.1980. (HOLOTYPE, BAFC 25525). Misiones: Puerto Libertad, Alto Parana
plantation, leg. Deschamps, 21.XII.1979 (BAFC 24535); Parque Nacional
Iguazu, leg. Bazzalo, 3. 111-1980... (BAPE. 25056) - BRAZIL: Manaos, banks
of Rio Tapajos,leg. Alves da Sousa, 9.X.1977 (BAFC 24429); ibid. ,Parque
Nacional Pedreiras, leg. ipse, 1977 (BAFC 25527).
The basidiomes have the external appearance of G. Luetdum
s..str. but the micromorphologic @ifferences are striking.
much smaller spores which are broadly ellipsoid, of the
"smooth" type, light context and frequent gasterospores both
in the context and dissepiments. The study of a part of the
holotype (S) of G. subambotnense P. Henn. showed that it
possessed gasterospores (not mentioned in the diagnosis), of
the same size and shape but differing in being ornamented
with veins anastomosing to form a sort of reticulum. Since
our materials only differ from G. subambotnense var. subam=
botnense in tne total lack of ornamentation of the gasteros-
pores and a slight variation in the spore measurements
which are 7-9 x 4-6,5 um in the holotype mentioned, we be-
lieve this warrants the new variety proposed.
Furthermore, the specimens studied appear to be close to
G. multtpltcatum (Mont.) Pat. var. vttaltz¢ Steyaert (1962),
which has also gasterospores but of a much larger size. We
have been unable to secure the holotype of the latter.
Figs. 23-25: basidiospores of the "rugose" type of Ganoderma luetdum
s. str. (BAFC 24425). 26-33: basidiospores of the "smooth" type of G.
restnaceum.26 and 28 from BAFC 24436; 27, 31-33: from BAFC 24431; 30:
from the holotype of G. sesstle (all 10000 x).
205
i
ji Ie
Ly)
—
if
a
Cultural features Figs. 72-82; 88-92; 94-101; 106.
Strains: BAFC n°247 = the HOLOTYPE (BAFC 25525). BAFC n°745 = Mi-
siones: Parque Nacional Iguazu (see BAFC 25056).
Codecnumbers) 123 3. 3." 10..134. 36. 37. Some aoe
Microscopié characters: growth fast, reaching 3-4,5 cm diam. in the
first week, covering the plates in 2 weeks; mycelial mat white (Pl. 1A
1); during the 3rd week ochraceous yellowish zones begin appearing ap-
pressed to agar (Pl. 12 B 7), at first irregularly at inoculum or at the
marginal area. Texture farinaceous during the lst week at the inoculum,
the rest felty, becoming after the 2nd week totally farinaceous (Fig.
106). Reverse only altered in the areas of coloured mycelium, becoming
yellowish to brownish. Margin regular, smooth. Odour sweetish. Oxidase
reaction: with tannic acid = ++++ with growth; with gallic:acid =+++++
without growth; with gum guaic= +.
Microscopic features: Marginal mycelium: formed solely by generative
hyphae during the Ist week, which are clamped, with scant branches,
septa restricted to clamps (Figs. 88-90), 2,6 um diam., staining with
phloxine or remaining hyaline; during the 2nd week: i) appear clamped
stag-horn hyphae that stain well with phloxine, of the same size as the
former, and with very abundant and thin ramifications which are very
characteristic, in general dichotomic, very intrincately branched (Fig.
92); ii) unclamped, fibrous, aseptate, hyaline, thick-walled slender
hyphae, 1-2 am diam., heavily branched (Figs. 77-78); iii) hyaline "cu-
ticular cells", originating from globose branches of hyaline generatives
(Figs. 72-74), attaining various shapes and sizes and arranged tightly
into a "pseudoparenchyma"; iv) dextrinoid chlamydospores, terminal or
intercalary, with a simple or double golden wall (Figs. 94-101), almost
spherical to ellipsoid, with dense contents that stain deeply with phlox-
ine, 11-18 x 9-15 um. Inoculum: with same features as the white mycelium.
Coloured mycelium: i) generatives as in i) above, scarce; ii) “cuticular”
cells with a thick golden wall arranged in a pseudoparenchyma which be-
comes compact and is responsible for the colour observed, shape and size
variable, each element ca. 9-47 um diam. (Figs. 75-76); iii) thick-wall-
ed clamped hyphae with golden walls, unbranched or with terminal irreg-
ular branches and unclamped sclerotized hyphae with narrow portions at
regular intervals, appearing as rosary beads, 2-6 um diam. (Figs. 79-82);
uy fibrous hyphae as in ii) above, very abundant, 2-4 um diam.; v) dex-
trinoid chlamydospores which may be terminal or intercalary, same as iv)
above. Submerged mycelium formed by: 1) generative hyphae with thin wall,
tortuous, much branched with short branches, staining with phloxine (Fig.
91), 2-5 um diam.; 2) chlamydospores as above; 3) "Cuticular cells" as
above. The submerged mycelium in the coloured zone have "cuticular
cells" with thick, golden walls and sclerotized generative hyphae, 2-8
am diam., as well as typical chlamydospores; generative hyphae are
scant.
This species has apparently not been described in culture before.
Figs. 34-45. Basidiospores of the "semirugose" type of Ganoderma
oerstedit. 34-35, 37-38: from BAFC 24441; 36, 41-42: from the holotype
of Ganoderma tuberculosum; 39-40, 43-45: from BAFC 24410 (all 10000 x).
307
GANODERMA RESINACEUM (Boud.)Pat., Bull. Soc. Mycol. Fr.
Siar Aa ets tee.
=G. chaffangeont Pat., ibid. 5: 74. 1889 (Fide Steyaert). = F.
restnaceus (Boud) Sacc., Syll. Fung. 9: 179. 1891. = G. sesstle
Murr.,. Bull. Torrey bot. Cl. 29: 604. 1902; North Amer. fie 9.
120. 1908 (NY!). = G. polychromum (Copel.) Murr., North Amer. FI.
9: 119219082 = G. pulverulentum Mury=,-. VWb1d.99: siz 190e
(NY!) =. = (4G: praelongum Murr., “ibid. .9P 121241908. (NYS). eee
subinerustatun Murr., ibid. 9: 120. 1908 (NY!). = G. argtlla-
eeum Murr. , ibids.9: 122. 1908 (NY!).” = Gy seezaune Mure.
ibid. 9: 123. 1908 (NY!). = G. subperforatum Atk., Bot. Gaz. 46:
337. 1908. = G. platense Speg., Bol. Acad. Nac. Cienc. Cordoba
2828363, 1926 (LPS!)
Figs: -7-103.26-33;" (60,
Annual, lignicolous, generally dimidiate to reniform,
sometimes circular, spathulate or ungulate; isolated or imb-
ricate, sometimes several pilei laterally confluent with
their pseudostipes free or fused into one; sessile, substip-
tate; 3.5-55 x 3-18 x 1-8 cm. Pileus Surface appianeter
concave or more or less infundibuliform, smooth, irregularly
rugose, concentrically sulcate and radially rugose or strong
ly tuberculose, laccate, brilliant or dull, sometimes dull
due to a thick deposit of spores. Central zone yellowish
brown, very light in young specimens (Pl. 12 H 12), darkening
with age from the centre towards the margin, with a broad
cream coloured to yellowish marginal band (Pl. 9 E 4)3 in
mature specimens there is no such gradation, the surface be-
ing dark reddish brown (Pl. 7 L 11) or light reddish brown,
homogeneous (Figs. 8,10). Margin sterile, thick, bDlune.
straight, incurved or recurved, yellowish cream in actively
growing specimens, dark reddish brown in mature ones. Pseudo
stipes 'Tateral or central,.very Short to long. silendemece
thick, blacky: Taccate, brilliant, 4-7 cm long. 2-5 chawide,
sometimes rudimentary. Section thin to very thick, thicken-
ing towards the base, 1-4 cm at about half the radius. Cont-
ext corky and soft or woody and hard; 2-5 cm thick, uniform-
ly brown (Pl. 13 H 9), or with a thin darker band above the
tubes. Hymenophore poroid, white to yellowish, becoming dark
brown when old (Figs. 7, 9) with a layer of tubes often dec-
urrent on the stem, 5-15 mm long, slightly lighter than the
context; pores circular, large to medium sized, 2-5 per mm,
89-309 um diam., dissepiments 27-267 um wide. Hymenium com-
posed only of basidia that are globose, soon collapsing, /-
14 x 8-20 um, 4-spored (Fig. 60). Basidiospores of the
"smooth" type, 9-13 x 5-8 um, with a thick endosporium,
light yellowish, ellipsoid, with numerous slender endosporic
pillars that. do not influence -the perisporiums which ers
smooth and thin, thus appearing "smooth" when observed with
the 0. M. at 400 x (Figs. 26-33). Dermis of the "hymenoder-
mis" type, composed of claviform elements with scant lumen
509
and blunt ends, originating from skeletal hyphae and arrang-
ed as in a hymenium, 7-16 um diam., dermis 13-43 um thick
(Fig. 62). Hyphal system trimitic, with clamped, thin-walled
generatives, with septa restricted to clamps, 1-6 um diam.,
sparsely branched, abundant. at the growth margin of the pil-
eus and dissepiments (Fig. 71). Skeletals of the "“arbori form
type, clampless, aseptate, with a thick, golden wall, with
few branches limited to the distal end, 3-8 um diam., very
long, forming the bulk of the context and dissepiments (Fig.
68). Binding hyphae of the "Bovista" type, clampless, asept-
ate, of limited growth, thick-walled, in general thinner and
paler than the skeletals, much branched, 1-4 um diam. (Figs.
69-70), only present in the context, as a rule very scarce.
Hosts: on dead trunks of Tzpuana ttpu (dead as a result of the
fungal attack); Quercus suber and undetermined hardwoods; at the base
Of Casuarina cunntnghamiana, Platanus acertfolta, Ulmus procera, Acucta
sp., Saltx, Prosopts algarrobilla, Blepharocalyx tweedtt and Robinta
pseudoacacta; also on stumps and roots of undetermined hardwoods. Do-
manski et al. (1967) record it also on Fagus and Alnus, although rarely.
Distribution: SOUTH AMERICA: Argentina (Buenos Aires, Capital Fede-
ral, Cordoba, Corrientes, Misiones, Tucuman and Salta); Uruguay; Vene-
zuela. NORTH AMERICA: U. S. (Connecticut to Missouri, Alabama, Louisia-
na). CENTRAL AMERICA: Cuba, Honduras, Jamaica. It has also been record-
ed from Euro-Asia and Africa (Central). It seems to have a world-wide
distribution in temperate and tropical areas.
Holotype: Fomes restnaceus Boud. (PC!).
Material studied: ARGENTINA: Buenos Aires: Parque Pereyra Iraola,
leg. Merlo, 15.1V.1969 (BAFC 24458); Llavallol, Santa Catalina, leg.
Deschamps et al., 4.11.1973 (BAFC 24460); Castelar, leg. Quiroga, 1.V.
1979 (BAFC24450); San Fernando, leg. Doyle (BAFC 78532). Hway 2, km 100,
leg. Wright & Deschamps, 24.IV.1971 (BAFC 25535) ;Martinez, leg. Wright,
21.11.1971 (BAFC 24461); San Miguel, Quinta Zemborain, leg. Campi, 22.
IV.1944 (BAFC 25529);La Plata, leg. Spegazzini, 8.V.1909 (LPS 24880)
Acassuso, leg. Soriano, 23.111.1949 (LPS 31293, sub G. Zuctdum); no
locality, leg. Spegazzini, II-1918 (LPS 24868, sub G. lorentatanum) ;no
data (LPS 24855). Capital Federal: Fac. Agronomia, leg. Maluh & Bargie-
la, 4-1-1979 (BAFC 24427); Golf Municipal, leg. Wright, 15.11.1968
(BAFC 24439); Villa Pueyrredon, leg. Romero, 16.11.1978 (BAFC 24440);
Jardin Botanico, leg. Agullo, 27.1V.1978 (BAFC 24412); Jardin Zooldgico,
leg. Astort, 29.1V.1979 (BAFC 24445); leg. Deschamps & Rovetta, 16.III.
1972 (BAFC 24459). Cordoba: Alta Gracia, leg. Spegazzini, 1.1925 (LPS
30989). Corrientes: 12 km NE of Curuzd Cuatia, leg. Singer, 8.11.1964
(BAFC 24436). Entre Rios: Concepcidén del Uruguay, La Salamanca, leg.
Hawryszko, ile XII.1961 (BAFC 24407); Colon, Parque Nac. El Palmar, La
Calera, leg. Del Busto & Deschamps, 2.1V.1971 (BAFC 24430); Dept°Rosario
del Tala, Palacio San José, leg. ipse, 9.1V.1971 (BAFC 24432); ibid.,
leg. Deschamps, 15.11.1972 (BAFC 25533); ibid, leg. ipse, 31.XII.1971
(BAFC 24434); Gualeguay, banks of Gualeguay River, leg. Wright, 27.1.
310
1951 (BAFC 24438); Salto Grande, leg. Deschamps, 31.X11.1971 (BAFC
24454); Gualeguaychu, Rincodn de Lauda, leg. Tonni, 21.VII.1974 (BAFC
24457); ibid., Parque Unzué, leg. Deschamps, 25.XI.1978 (BAFC 25530);
Arroyo Isletas, leg. Bettucci, 12.XI1.1963 (BAFC 24431); no data (BAFC
24453). Misiones: Colonia Belgrano, leg. Gomez, 11.1965 (BAFC 24420).
Salta: Alemania, leg. Bettucci & Guerrero, 29.1.1965 (BAFC 24409),
Tucuman: garden of Lillo Inst., leg. Wright, 27.11.1971 (BAFC 24411);
San Javier, leg. Rosa Mato n°2186 (LPS 26190 ex MVM); no data (LPS
24866). CUBA: Prov. Santiago, Alto Cedro, leg. Earle & Murrill 536,
19/20. 111.1905 (HOLOTYPE of G. praelongum Murr., NY); Prov. Habana:
pr. Santiago de las Vegas, leg. Earle 658, 5.VII.1904 (HOLOTYPE of a.
argillaceum Murr., NY). UNITED STATES: New York, Bedford Park, V-1902
(HOLOTYPE of G. sesstle Murr., NY). BRITISH HONDURAS: Puerto Sierra,
Rio Esperanza, leg. Wilson 607, 28.11.1903 (HOLOTYPE of G. ntt¢dwn Murr.
NY). GRENADA: leg. Broadway, 4.X1.1905 (HOLOTYPE of G. puZverulentun
Murr., NY). JAMAICA: Hope Gardens, leg. Earle 176, 26.X.1902 (HOLO-
TYPE of G. subtnerustatum Murr., NY). URUGUAY: Dept°Canelones, Parque
Nac. Carrasco, leg. Garcia Zorr6én, 16.V~1960 (MVHC 2473).
Steyaert (1972) studied the holotypes of G. sessile, G.
praelongum, G. argtllaceum, G. restnaceum, G. polychromum,
G. subperforatum and G. chaffangeontt, and found that they
all possessed identical micromorphological features, partic-
ularly the "smooth" type of spores, for which reason he cor
idered them synonyms, the valid name of the species being
Ganoderma restnaceum (Boud.) Pat. We have been able to
study all the holotypes of Murrill's species deposited at
NY and we agree with him, but found that to the above list
must be added G. ntttdum and G. pulverulentum, as well as
G. platense Speg., the holotype of which was studied in det-
ait) CEP San
The description of G. subinerustatum given in North Amer-
ican Flora by Murrill, does not agree with the holotype
(Murrill, 1908: 122), since the spores are given as 8 x 4
Am, whereas Our measurements are 9-11 x 6-8 um. In the diag-
nosis of G. ntttdum Murr., in the same paper, no mention is
made of the spores; its holotype, according to our measure -
ments, has spores measuring 9-13 x 5-7 wm.
Spegazzini (1926) reported from Argentina G. lorentztanum
Kalchbr., including stemless, pleuropodal and centrally
stipitate specimens, a concave, more or less infundibuliform
pileus, and ovate, smooth to slightly ornamented spores
measuring 10-12 x 5-7 ym; at first we believed they were mere forms
of G. restnaceum, but a detailed study of them showed there
were really significative differences between those specim-
ebs and G. restnaceum. We do not know whether Spegazzini
Studied authentic materials of G. lorentztanum Kalchbr.,and
we were unable to locate the holotype in order to solve the
Figs. 46-50. Smooth type basidiospores of Ganoderma subambotnense
var. Laevtsporum 46,49: from BAFC 24535; 47: from BAFC 25056; 48:
from BAFC 25527; 50: from BAFC 25525. 51-56. Semirugose type of bas-
idiospores of G. gonatum. 51-52: from the Holotype; 53,55: from BAFC
24416; 54: from BAFC 24449; 56: from BAFC 24414 (fig. 51, 9000 x; the
rest are all 10000 x).
Dae
46-50. 52-56
eee ilies
2p
SES
problem.On the other hand of two other specimens identified
as G. lorentztanum Dy spegazzini at LPS; one has “smooth®
type of spores and the other "semirugose" type. Thus, the
former must be included in G. restnaceum and the latter in
G. tubereulosum.
The study of the holotype of G. lorentztanum Kalchbr.
would be important to ellucidate the cospecificity of this
Species with G. restnaceum, Since should they prove to be
the same, G. lorentztanum would be the valid name.
Cultural foearures Figs. 75-78; 88-93; 96-103.
Strains: BAFC n%2221 = ARGENTINA: Buenos Aires, Llavallol (See BAFC
24460). BAFC n°2775 = Capital Federal, Villa Pueyrredon (see BAFC
24440). BAFC n°2354 = ibid. (see BAFC 24459). BAFC n°658 = ibid.,
Jardin Botanico (see BAFC 24412). BAFC n°2318 = ibid., calle Julian
Alvarez y Araoz. BAFC n°2813 = Entre Rios, Arroyo Isletas (see BAFC
24431). BAFC n°228 = ibid., Dept°Rosario del Tala, Palacio San José
(see BAFC 24432). BAFC n°2294 = ibid. (see 25533). BAFC n°443 =
ibid., leg. Del Busto & Deschamps, 25.XI.1968. BAFC n°1009 = ibid.,
Gualeguay (see BAFC 24438). BAFC n°445 = ibid., Colén (see BAFC 24430).
BAFC n°2576 = Tucuman: garden of the Lillo Inst. (see BAFC 24411).
BAFC n°834 = CANADA, Ontario, Ottawa, 12.1X.1941 (DBFP 10222). BAFC n°
fiz = ex NY 5SI9T-sentrby Dre Robbins. 1BAFC. n° 716 ="ex.CBS 152222
Code number! 2.3.) 8. (10).°1345; 3652.37. (39% (42) 20 (47 Sa
Macroscopic characters: growth moderately rapid, covering Petri dish-
es in 2-3 weeks. Mycelial mat transparent white at first, later remain-
ing snow white with rather loose mycelium; in some cultures the inocul-
um becomes cream coloured with a few light yellowish zones appressed
to the agar, irregularly distributed. Texture farinaceous during the
first week at the inoculum, the rest subfelty, later the whole surface
becoming farinaceous and the mycelium much denser (Fig. 102); in other
cultures there is an alternation of dense, fan-shaped areas, with much
less dense mycelium (Fig. 103), or zones that show alternation of dense
and thin mycelium. Reverse: only altered in correspondence with zones
of coloured mycelium, which becomes brown. Margin regular, smooth,with
mycelium appressed to agar. Odour: slightly fungic to somewhat sweetish.
Oxidase reaction: with tannic acid = ++++ with or without growth; with
gallic acid = +++ or ++++ with or without growth; with gum guaiac = +.
Microscopic features: marginal mycelium: composed during the lst
week only of generatives, staining or not with phloxine, clamped, thin-
walled with septa restricted to clamps, sparsely or not branched (Figs.
88-90), 2-6 am diam. From the lst to the 3rd weeks the following appear:
i) "stag-horn" generatives, staining with phloxine, whose ends begin to
branch out in numerous and very thin ramifications, mostly dichotomic,
that become intertwined and form a dense reticulum (Fig. 92), 2-4 um
Fig. 57 .Aspect of hymenium, basidiospores and gasterospores present
in the dissepiments of G. subambotnense var. laevisporum. Fig. 58. As-
pect of hymenium and basidiospores of G. oerstedit. Fig. 59. Aspect of
hymenium and basidiospores of G. zonatum. Fig. 60. Aspect of hymenium
and basidiospores of G. restnaceum. Fig. 61. Aspect of hymenium and
basidiospores of G. Zuetdum. (All figures 1500 x).
51S
58
—p SS SS
Sock
H PyAg
sD 7p sea | UE ye
S : SES
314
diam;ii) chlamydospores both intercalary and terminal, abundant, the
first originating from generatives, the latter from the ends of stained
generatives; thick-walled, golden, with dense contents, ellipsoid, sper
ical or ovoid, dextrinoid, 9-22 x 7-18 um (Figs. 96-101); iii) fibrous
hyphae that are clampless, aseptate, thick-walled, hyaline or pale, not
Staining, heavily branched, in some cases forming the bulk of the mycel-
jum, 1-4 um diam (Fig. 77-78); from the 3rd to 4th weeks, there appear
“cuticular cells", hyaline, originating from globose ramifications of
generatives, of very irregular shape and size, 10-30 um each, much ap-
pressed and forming a pseudoparenchyma that may become very compact
(Fig. 93). Inoculum: same features as the marginal mycelium. Yellowish
zones: formed by the same elements as the rest of the mat but the "cut-
icular cells" have a thickened golden wall (Figs.75-76); in a few cases
sclerotized clamped hyphae appear, with thick, golden walls, but are
not abundant. Submerged mycelium composed of tortuous generative hyphae
with thin walls, that may stain or not, and have short but numerous
projections, 1-5 am diam. (Fig. 91). Also chlamydospores similar to
those of the margin and a few "cuticular cells" may be found.
Observations: Culture BAFC n°2354 was the only one that
fruited in the Petri dishes after the 5th week, in the form
of an elevated mound in which a great density of fibrous
hyphae concentrated; on them the pores were formed with fert-
ile hymenium composed of globose basidia and basidiospores
of the "smooth" type, 8-ll x 5-6 am.
There exists a difference that allows the separation of
the strains in two obvious groups, while the remaining feat-
ures are similar in all cases, namely, the presence or abs-
ence of “cuticular” celis.sStrains, BARC. no 2o0-Aoeuc 7. meen
1009 always formed these cells. The rest did not. The basid-
jomes corresponding to these cultures did not exhibit any
differences.
The cultural characteristics of the strains studied, coin-
cide with those given by Nobles (1948), excepting the "cutic-
ular cells, which may or may not be present, and the reverse
that darkens but does not appear to become discoloured, as
mentioned by her. Concerning this aspect, Stalpers (1978)
states that it discolours locally and becomes locally colour-
ed, with which observation we agree. Cultures from the CBS —
and NY identified as G. restnaceum were studied and they also
didn norushows +cutacutar cel ls
GANODERMA OFRSTEDII (Fr.)Torrend, Broteria 17: 37. 1920.
=Fomes oerstedit Fr., Nov. Symb. Myc. p. 63. 1855 (S!). =G. tu-
berculosum Murrill, North Amer. Fl. 9 (2): 123. 1908 (NY!).
Figs. 13-16; 34-45; 58,
Annual, lignicolous, dimidiate to reniform, isolated, cometimes imb-
ricate, sessile or stipitate, small to large, 10-40 x 5-25 x 2-10 cm.
Figs. 62-71. G. subamboinense var. laevtsporum. 62: hymenodermis ;
63-66: gasterospores from the context and dissepiments; 67: aspect of
context; 68: skeletal hyphae; 69-70: binding hyphae; 71: generative hyp-
hae. Figs 72-78: cultural features (continued next page)
316
Pileus surface radially rugose and concentrically sulcate or
strongly tuberculate, frequently umbonate behind, laccate,
brilliant,but mostly appearing somewhat opaque due to the
deposition: of a thick layer of brown:<spores (Figs. 14) ite).
central zone dark reddish brown (Pl. 56 L 12), almost black,
to lightareddish brown (P1i 6 lb. 12),emuUch Tighter: towards
the margin; generally the young specimens exhibit a wide
creamish yellow marginal band (Pl. 9 E 4) that becomes homo-
geneous dark in mature ones. Margin sterile, thin to thick,
acute to blunt, smooth or wavy, white to cream-colour (PI.
9 B 1) in actively growing specimens, then straight, reddish
brown in adult specimens, then incurved. Stem dark reddish
brown, laccate,. brilliant, thin’ and cylindric? to® thickeanc
depressed, 5,5-9 scm: Tong,. 1. 1-5’ cm ‘widex *Section, tninero
thick, 1,5-5 cm deep at about half the’ radius, thickengng
towards the base. Cutis: thin,, black brilliant Contextycorky,
brown (P1 13 E 8), with a thin darker band over the tubes,
or homogeneous: dark brown, (P1.97. € 11); sup om seemtiitcr.
Dermis of the "hymenodermis" type, composed of claviform
tips of skeletal hyphae arranged like a hymenium, lumen small
and blunt ends, 6-14 um diam., 17-40 um wide (Fig. 62). Hym-
enophore poroid, greyish white to yellowish (Pl. 9 J 5) dark
ening sometimes with age; pores small, circular to somewhat
irregular, 4-6 per mm, 70-270 am wide (Figs. 13, 15); tube
layer: slightly lighter than context, up to 43) cm thick weqice
sepiments 27-180 um thick. Hymenium composed only of scant
globose, 4-spored basidia, 9-16 x 9-10 um (Fig. 58). Basid-
iospores of the "semirugose" type, brown in mass, broadly
ellipsoid to almost ovoid (Figs. 34-45), with truncate ape,
hyaline perisporium smooth and thin, endosporium thick, gold-
en brown, with numerous endosporic pillars reaching the per-
isporium, rumpling it slightly and making it appear slightly
rugose, 9-14 x 6-9 am. Hyphal system trimitic with clamped,
thin-walled generatives with septa restricted to clamps, un-
branched, 2-4 um diam., almost always restricted to dissep-
iments (Fig. 71).Skeletals of the "arboriform" type, clamp-
less, aseptate, with a thick, golden wall,subsolid, with a
72-74: formation of “cuticular cells" from globose branches of generat-
ive hyaline hyphae; 75-76: thick-walled "cuticular cells"; 77-78: fib-
rous hyphae.
Figs. 79-93. Cultural features of G. subambotnense var. laevtsporum
and of G. luetdum. 79-82: Slerotized hyphae which are identical in both
species, in 79 and 81 with thickenings that make them appear like rosary
beads; in 80 only in the former species, with irregular branches and
also unbranched. Figs. 83-87. G. oerstedit. Cultural features. Sclerot-
ized hyphae with 83: short branches, 84: with numerous branches in all
planes, 85-87: with claviform ends; 88-91: thin-walled generative hyph-
ae,) 88-90 from aerial mycelium; 91: from submerged mycelium; 92: "stag-
horn" hyphae; 93: formation of hyaline "cuticular cells". Figs. 94-101:
chlamydospores. 94-95: double-walled chlamydospores of G. subambotnense
var laevtsporum. 96-97: simple-walled intercalary chlamydospores; 98-
101: terminal ones (all figures 1500 x).
Sst
few restricted branches at the apical end (Fig. 68). Binding
hyphae aseptate, unclamped, with limited growth, heavily
branched, thick-walled, generally thinner and paler than
Eerie 1-4 um diam., only present in the context (Figs.
69-70).
Hosts: on stumps of undetermined hardwoods, fallen trunks, dead
Quercus suber, Pinus spp., Ocotea acuttfolta, Casuarina cunninghamtana,
Sptraea cantontensts, Scutta buxtfolta and Lonchoecarpus sp.
Distribution: ARGENTINA (Buenos Aires, Cordoba, Corrientes, Entre
Rios, Misiones and Tucuman)y Murrill (1915) records it on dead wood in
British Honduras, Panama and Western Jamaica; Dennis (1970) records it
for Guyana.
Material studied: ARGENTINA: Buenos Aires: Acassuso, leg. Sauval,
17.1IV.1980 (BAFC 25528); Punta Lara, leg. Wright, 21.11.1971 (BAFC 2441
24423); Gral Alvear, leg. Di Riccio, 10.1V.1968 (BAFC 24410, 24411) .
Cordoba: Salsipuedes, leg. Godeas, 5-III-1973 (BAFC 24442). Corrientes:
Saladas, Paso Naranjito, marginal forest of Rio Santa Lucia, leg. Ro-
vetta et al., 12.VII.1973 (BAFC 24413); Mburucuya, Estancia Santa Tere-
sa, leg. Wright et al., 16.VIII.1972 (BAFC 24424); Dept°Monte Caseros,
Rio Uruguay, Isla Itacumbd, E bank, leg. Irigoyen, 20.XII.1977 (CTES
438). Entre Rios: Salto Grande, leg. Deschamps, 31.XI1.1971 (BAFC 24426).
Misiones: Parque Nacional Iguazu, Cataratas, leg. Wright et al.,25-X-
1973 (BAFC 24437). Tucuman: no data (LPS 24866 as G. lorentztanum);
no data (LPS 24869); no data (LPS 2497 as G. orbtforme). BRITISH HONDU-
RAS: leg. M. Peck (HOLOTYPE of G. tubereulosuwm Murrill, NY). COSTA RICA:
leg. Oersted (HOLOTYPE of Fomes oerstedit Fr., S; part at BPI).
Observations: Our materials coincide with the features
of the holotype in everything except that their spores are
somewhat more polymorphic, varying from ellipsoid to broad-
ly ellipsoid. Some specimens resemble macromorhologically
G, resinaceum (Boud.)Pat., but can be readily distinguished
from this species by the basidisopores that are of the
"smooth" type, and the number of pores per mm. This is the
first record for Argentina. Steyaert (1980) places this
Species in his Section Characodermaon the basis of the thinner
hyphae ending in a sphaeroid, which form the hymenodermis.
We believe this feature is sufficiently variable to invalid-
ate this proposal, precisely because in the present species
such character, aS Steyaert (loc. cit.) nhimselt staves are
intermediate in nature.
Cultural features Figs. 83-92; 104-105.
Strains: BAFC n°218 = ARGENTINA: Buenos Aires: Gral Alvear (see
BAFC 24411). BAFC n°178 = ibid. (BAFC 24410). BAFC n°2249 = Cordoba:
Figs. 102-103. Cultural features of G. restnaceum. 102: strain BAFC
n°2576; 103:strain BAFC n°113. Figs. 104-105. Cultural features of
G. oerstedit. 104: strain BAFC n°218; 105: strain BAFC n°2249. Fig. .106:
Cultural features of G. subambotnense var. laevtsporum. Fig. 107: Cult-
ural features of G. Zuctdum s. str. strain BAFC n°112 (all figures i11-
ustrate 3 week old cultures).
o19
320
Sree (BAFC 24442). BAFC n°2382 = Corrientes: Saladas (BAFC
24413).
Gode number?) (2. 3.8. 10032. (34). 36.372) 39: 42. (43) abs haeeoe
Macroscopic characters: growth rapid, covering Petri dishes in 2-3
weeks. Mycelial mat subfelty and white, at first transparent, later be-
coming totally farinaceous with yellowish zones (Pl. 13 K 5), irregular-
ly arranged on the inoculum and the margin opposing it (Fig. 105) ,which
may reach a large surface, or closely packed, felty at the inoculum,
borders and discrete mounds irregularly scattered, somewhat elevated,
later becoming confluent with ochraceous yellowish zones (Pl. 9 K 5),
irregularly appearing on a dense mycelium (Fig. 104). Odour sweetish.
Reverse altered in correspondence with the areas of coloured mycelium,
becoming light to dark brown. Oxidase reaction: with tannic acid = ++++
with or without growth; with gallic acid = +++ with or without growth;
with gum guaiac = +.
Microscopic features: Marginal mycelium during the lst week composed
mainly of generative, clamped hyphae with septa restricted to clamps,
thin-walled, scarcely or not at all branched; in general, the branches
originate at the clamps staining with phloxine, or hyaline, 1-4 wm
diam. (Figs. 88-90); during the lst or 2nd week, "stag-horn" hyphae
appear, clamped, staining well, with dichotomically branched tips, dev-
eloping numerous and very thin ramifications (less than 1 um diam.),
very intrincate and intertwined (Fig. 92); fibrous hyphae clampless,
aseptate, hyaline, with a pallid thick-wall, heavily branched, 1-2 um
diam., these appear mostly the 2nd week, and when the margin becomes
dense these constitute the bulk of the mycelium; otherwise they are not
so numerous (Figs. 77-78); intercalary chlamydospores arising along
clamped generativess, spherical to ellipsoid, with dense contents,
8-18 x 5-11 um, very abundant or totally absent (Figs. 96-97). White
farinaceous mycelium is composed of: i) generatives like those of margin,
scarce; ii) "cuticular cells", hyaline, originating from globose ramif-
ications of hyaline generatives (Figs. 72-74), very abundant, arranged
in an appressed pseudoparenchyma, of various shapes, globose to very
irregular; each element 4-30 um diam.; iii) chlamydospores as those des-
cribed above. Yellow farinaceous mycelium: with same elements as former,
but the "cuticular cells" have golden, thick walls (Figs. 75-76); gen-
eratives with sclerotized golden walls are also present, which stain.
well with phloxine, and bear short branches, 1-4 um diam.(Fig. 83). Zone
of dense mycelium and elevated mounds: formed mainly by fibrous, abund-
ant hyphae, with same features as those of margin; "cuticular cells"
also present, mainly hyaline, as well as stag-horn hyphae. The zones of
dense mycelium that become yellowish are formed by sclerotized clamped
hyphae or not, sometimes very abundant, with a golden thick wall, scant
lumen and claviform ends similar to the elements of the dermis of the
basidiome, or numerous small branchlets branching in all planes, 2-9
um diam. (Figs. 84-87). Submerged mycelium: formed by the same elements
as the marginal, but generatives are more tortuous and possess numerous
short branchlets (Fig. 91).
This species has not been described in culture before.
Numerous differences are observed between strains 178 and
218, and between strains 2383 and 2249. The first two exhib
it a tight felty texture (Fig. 104) complete absence of
chlamydospores, abundance of fibrous hyphae and less proport
10n, of “cuticular cells"; the latter two present;a farinec-
SZ.
@mustexture (Fig. 105), presence of intercalary, dextrinoid
chlamydospores, smaller proportion of fibrous hyphae, and
compact pseudoparenchyma of abundant "cuticular cells". Co-
incident with these differences, the basidiomes from which
strains 218 and 178 were obtained, have larger pores than
the remaining specimens studied, the average being 4 pores
per mm. Until further studies on interfertility can be car -
ied out that could prove whether these differences are sign-
ificant, it is preferable to maintain these materials as G.
oerstedtt.
GANODERMANZONATUM Muri), Bull. Torrey. bot. -cl. -29% 606.
IOOZ enon huAneY. FIs 9.42) 2120261908,
= G. sulcatum Murr., ibid. 29: 607. 1902; North Amer. Fl. 9 (2):
120. 1908 (NY).
Bigs.) 1/=22 590 1-56:." 59,
Annual, isolated, dimidiate and laterally stipitate, or
several pilei imbricate on a common stem, 6-20 x 4,5-20 x
0,5-5 cm. Pileus surface smooth or concentrically sulcate,
usually with 1-3 furrows, the inner one usually deeper, thus
exhibiting a wavy appearance, sometimes also radially rugose;
liaiccatve, brilliant, central zone reddish brown (P17 I 11)
becoming lighter towards the margin , with a narrow orange
brownish marginal band (Pl. 12 D 11), often delimited by the
external furrow and the margin (Figs. 17, 19-20). Margin
Slewelese D hunts strarght. yellowish whitish <@P1.° 11 °B 1).
Stem iconcolorous with pileusy laccate.-brilliant,. 3,5-5 cm
long, 2-4 cm wide, vertical,cylindric or depressed. Section
8-10 mm deep at about half the radius, thicker at the base.
Cut saenriinie DIack sbi liante.tcontext. brown Pde 14061206
Slightly lighter at the surface of pileus, 3-44 mm thick,
corky, soft. Dermis of the "hymenodermis"type, composed of
the claviform ends of skeletal hyphae arranged in a palisade-
like hymenium, 6-11 um diam., with small lumina, blunt ends
anGstnick, colden walls 3-dermis 1/-33 um-thick (Fig. 62).
Hymenophore poroid, tube layer 1-5,5 mm long, slightly light-
er than context, (Pl. 13 C8), decurrent on stem; concolorous
with margin, pores 3-5 per mm, 98-267 wm diam., circular to
Subangular, greyish to slightly brownish upon maturation
(Figs. 18, 21-22); dissepiments 30-89 am. Hymenium not per-
sistent, composed only of scant basidia, 4-spored, 11-17 x
7-11 um (Fig. 59). Basidiospores of the "semirugose" type,
very characteristic, ellipsoid, long and slender, with trunc-
ate apex or, more frequently, rounded, 11-14 x 5-7 um, peris-
porium hyaline, smooth and thin, endosporic pillars numerous,
reaching the perisporium and slightly rumpling it, thus mak-
ing it appear slightly rugose; endospore golden and thick
(Figs. 51-56). Hyphal system trimitic, with hyaline, thin-
walled, clamped generatives, with septa restricted to clamps,
Sparsely or not at all branched, 2-4 um diam., present in the
growth margin of pileus and dissepiments (Fig. 71). Skeletals
of tne “arboriform’ ‘type, long, with a thick.,, golden walt,
clampless, aseptate, with 2-3 branches of same diam. as
mother hyphae at distal end (Fig. 68), 3-6 um diam., the
most abundant in the context and dissepiments. Binding hyph-
ae of the "Bovista" type, clampless, aseptate, of limited
growth, heavily branched, generally thinner and lighter than
Skeletals, 1-3 um diam., thick-walled, restricted to the con
textiGbigs 69 =7.0)
Hosts: on dead stem of palm (Butta yatay) and dead wood and roots
of living Tzpuana ttpu.
Distribution: U. S. (Florida, Georgia); ARGENTINA (Corrientes and
Entre Rios ).
Material studied: ARGENTINA: Corrientes: Dept°Cosme, Paso de la Pa-
tria, leg. Singer, 5.1V.1957 (BAFC 24416); Mburucuya, Estancia Santa
Teresa, leg. Pedersen, 29.II11.1977 (BAFC 24414). Entre Rios: Parque
Nacional El Palmar, leg. Mercuri, 25.11.1979 (BAFC 24449). UNITED STATES:
Florida leg. Underwood, 1914 (HOLOTYPE , NY!); ibid., leg. Lloyd, 1.1897
(HOLOTYPE of G.suZeatum Murr., NY!). CUBA: leg. Murrill (BAFC 25601, as
G. tuberculosum).
It was not possible to obtain cultures of this species.
According to Murrill, the only differences hetween G. zo-
natum and G. suleatum lie in the pileus surface. Both holo-
types were studied in detail, their microscopic features be-
ing identical, especially the basidiospores, which are long,
Slender, ellipsoid, "“semirugose" and of the same size. Pores
are identical. All this coincides with the Argentine specim-
ens(their spores being very slightly more rugose). We do not
agree with Murrill's description in North American Flora,
Since our measurements of the spores from the holotypes are
11-14 x 5-7 am and not 8-10 x 4-6 um as he states, and the
pores are 4-5 per mm and not 3-4.
We agree with Steyaert and Overholts that both species are
Synonyms. The former revised the holotypes in 1962 establish
ed their identity, and the latter published them as synonyms
but- as ‘varieties of P. ‘luctdus Leys....:Fre, under “the epi tier
~. Luetdus var. zonatum (Murr. )Overh. Only interfertility
Studies will be able to prove whether or not this taxon can
be separated from G. lLuctdum. For the time being we prefer
to consider it asaseparaite:.
DISCUSSION
According to our studies, the G. Zuetdum-complex would
thus appear to be represented in Argentina by five species,
namely, G. Luctdum s. str., G. resinaceum (Boud. )Rat.amc, ec
natum Murr., G. oerstedtt (Fr.)Torrend, and G. subambotnen-
se P. Henn. var.laevtsporum var. nov. Spegazzini (1926) re-
corded other species for Argentina, which we have been undle
to find in his herbarium, namely G. eupreum, G. sceleton and
G. fornicatum. G. platense Speg., whose holotype was stud-
ied, resulted another synonym for G. restnaceum. Additional
Synonyms for this species proved to be G. pulverulentum, G.
ntttdumandG. subtnerustatum.,. The question of the validity
Of G. multtplicatum var.vitalti Steyaert, G. lorentatanum
ig 50)
Kalchbr., and G. orbtforme Fr. remains pending a thorough
study of their respective holotypes.
Since other holotypes and authentic materials of species
that have as yet not been found in Argentina were studied,
but were recorded from other South American countries, a
preliminary general key for the complex has been included
in this paper.
The type of dermis is useless for the separation of spec-
ies, but permits the separation of "complexes" of species,
such as in the present case. In this we agree with Steyaert
(1972, 1980) and Imazeki (1939); they considered the exist-
ence of three or four (Steyaert, 1980) subgenera on this
basis. Regarding the hyphal system of basidiome construct-
ion, our results confirm those obtained by Hansen (1958).
Cultural studies did not exhibit significative differen-
ces and ought to be used with caution. We attempted several
methods for germinating spores with the hope of obtaining
monosporous cultures in order to verify interfertility pat-
terns, but were unsuccessful. This had been done previously
by Merrill (1970) with similar results. We also tried the
methods proposed by Aoshima (1953) and Brown (1970) for
spores Ot G. applanatum,out also failed. We did not test
Lim's method (1970). Equally unsuccessful were our attempts
to obtain monokaryotic mycelia from dikaryotic ones (dedi-
karyotization). More studies in this line are urgently need-
ed.
ACKNOWLEDGEMENTS
We wish to express our gratitude to the Instituto Forestal Nacional
(IFONA), of Argentina, for granting Miss Bazzalo a fellowship for this
Study; to the Curators of the herbaria of the New York Botanical Gard-
en, Jardin Botanique National de Belgique, Instituto "Carlos Spegazzi-
ni" and Instituto de Botanica del Nordeste (Corrientes) for the gener-
ous loan of holotypes and other specimens in their keeping; to Mr. Emi-
lio Del Busto, in charge of the culture collection of the Mycology Lab-
oratory, for his unflinching cooperation in our cultural work; to the
SEM Service of the Consejo Nacional de Investigaciones Cientificas y Téc-
nicas, for the work on spore ornamentation, and to the many colleagues
who have at different times and in different ways made this work poss-
ible. Special thanks are due to Dr. Leif Ryvarden (Oslo), Frances F.
Lombard and Michael Larsen (Madison) for critically reading the type-
script and making invaluable suggestions.
LITERATURE
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mycetes. ibid. 43: 1097-1139.
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gos Polyporaceos. Publ. Inst. bot. Sao Paulo. 22 p.
TORREND, C. 1920a. Les Polyporées du Brésil. I. Le genre Ganoderma.
Broteria (Bot .) 18: 28-43.
---------- 1920b. Les Polyporées du Brésil. II. Polyporacées stipitées.
ibid. 18: 121-142.
NOTE.
After this paper was ready for the printer, two works related to a
new species of the Ganoderma luctdum-cdmplex from Europe came to our
attention. Jahn, Kotlaba & Pouzar (Westfalische Pilzbriefe 9 (6): 97-
124, 1979/1980) described Ganoderma atkinsoni n. sp., very akin to G.
luctdum, but differing by «ts rugose, wider spores measuring (9,8)-10,4
-12,5 (-43,5) x (6,8)-7,3-7,8 (-8,5) um, often Larger carpophores, the
dank colour of the pileus and other substrates, mostly conifers. This
species had been named G. pseudoboletus (= G. Lluetdum) forma montanum
by Atkinson. Kotlaba & Pouzar (Ceska Mycologie 35 (3): 121-133. 1981)
added further data on this species, which seems restricted to Central
Europe and Great Britain.
MYCOTAXON
Vows SRV ING elope 826-75 5.0 October-December 1982
Es OE ae ds a ee a SRNR Se
TWO NEW SPECIES OF HELOTIALES FROM THE EASTERN HIMALAYAS
R. SHARMA
Botany Department, Panjab University, Chandigarh-160014, India
and
RICHARD P. KORF
Plant Pathology Herbarium, Cornell University, Ithaca, NY 14853 USA
During fungal forays undertaken by the senior author to the Eastern Hima-
layan regions of India and part of Bhutan, several interesting taxa be-
longing to the Helotiales have been collected. These include two new spe-
cies belonging to the genera Bisporella Sacc. and Crocicreas Fr. emend.
Carpenter, described and illustrated here.
Bisporella calycellinoides R. Sharma & Korf, sp. nov.
LEE GS ¢ welemercs
Apothecia gregaria, parva, sessiles, cupuliformes, cremea ad brunneola,
ad 1.5 mm diam., extus minute puberula, as basim annulo brunneo. Hymenium
dilutum aurantiacum, in sicco cyathiformis, peraurantiacum. Asci octospo-—
ri, poro iodo caerulescente, 58-65 x 6.5-7.5 pm, cylindracei-clavati,
e uncis natis. Ascosporae hyalinae, biseriatae, guttulatae, guttulae po-
lares, 9-11 x 2-3 pm, fusiformes, inaequilaterales. Paraphyses lanceola-
tae, septatae, leviter tumidae sub apice, ramosae, ad 6 pm latae, ascos
superantes ad 18 pm. Excipulum ectalum ex textura intricata ad 36 pm la-
tum, hyphae e cellulis parietibus incrassatis, cellulae plusminusve rec-—
tangularae. Excipulum medullatum ex textura intricata, ad 44 pm latum,
hyphae hyalinae, cellulae leptodermaticae, ad 2 pm diam. - Holotypus:
In foliis mortuis, Bhutan, Nawephu, Sept. 18, 1980; Raghunandan Sharma,
PAN 17518. In herbario universitatis Panjab Cryptogamarum, Chandigarh.
Apothecia gregarious, small, sessile, cupulate, hymenium light orange,
margin and external surface cream coloured, minutely downy, up to 1.5
mm diam, with a dark brown ring at the point of attachment, hymenium be-
coming deep orange, apothecia becoming more cupulate on drying. Asci 8-
spored, J+, 58-65 x 6.5-7.5 pm, cylindric-clavate, base slightly swollen,
apex round, arising from croziers. Ascospores hyaline, biguttulate, gut-
tules polar, 9-11 x 2-3 pm, fusoid, inequilateral, biseriate. Paraphyses
lanceolate, swollen a little below the apices, branched, septate, up to
6 pm broad, projecting up to 18 pm beyond the ascus tips. Excipulum dif-
ferentiated into two zones: ectal excipulum up to 36 pm thick, of thick-
walled, gelatinized, undulating hyphae, cells hyaline, 11 x 5.5 pm, some-
what rectangular; medullary excipulum of textura intricata, up to 44 pm
thick, hyphae hyaline, thin-walled, up to 2 pm broad. Hairs formed when
outermost cells are, drawn out into small, smooth, 1-septate, hyaline
hairs with obtuse tips, up to 18.5 x 3.5 pm; giving the apothecium 1ts
downy appearance. Hyaline hyphae near the base of the apothecium were
up to 2 pm in diam, and observed to penetrate the host.
ETYMOLOGY: Refers to the calycellinoid ring at the base of the apothecia.
HABITAT: On dead angiosperm leaves.
527
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ag
; 8,
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PNY
Y
F
N
a,
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Eee,
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fH
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SL
ee
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\ XK (Nl See AHI CE
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gh margin.
C. 8 ascospores.
Dix L000. (del. R.. Sharma)
FIG. 1. Bisporella calycellinoides. A. Transverse section throu
D. 3 paraphysis apices. A, B: x 500; C,
B. Transverse section through base of apothecium.
328
TYPE LOCALITY: Bhutan: Nawephu.
HOLOTYPE: PAN 17518, R. Sharma, September 18, 1980; ISOTYPE: CUP-IN 612.
NOTES: The undulating, thick-walled, gelatinized ectal tissues place this
species in Bisporella. The habit on leaves, and the distinct brown basal
ring immediately recall Calycellina, as does the yellow colour. In both
genera hair-like processes are formed in some species.
Crocicreas carpenteri R. Sharma & Korf, sp. nov. RIGS. 35.4]
Apothecia sparsa, parva, sessiles, nigra, in sicco porphyrea, cupulifor-—-
mes, ad 1 mm diam. Extus pilosus, pili saepe fasciculati, ad marginem
formans dentes, cylindracei, contracti versus apicem, atrobrunnei, multi-
septati (ad quatuordecimseptati), pachydermi, aspri granulis, ad 130 pm
longi, cellulae ad 11.5-3.5 pm, parietes cellularum ad 2 pm crassi. Hyme-
nium dilute brunneum. Asci octospori, poro jodo non caerulescente, 90-108
x 7.5-9 pm, apex rotundatis, incrassatus ad 2.0 pm. Ascosporae hyalinae,
multiseptatae (at maturatim usque ad septemseptatae), 15.5-21 x 2.5-3.5
FIGS. 2, 3. Transverse sections through margins, x 500. FIG. 2. Bisporel-
la calycellinoides. FIG. 3. Crocicreas carpenteri. (photos: R. Sharma)
( (id
SEP ape i IN |
= vy, Uy I
Coin <aitia ‘ai
Roar ie ot en Lis b
lore = ae, iff}
Apa YES
OES
WE KS Ye
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WAY
NG SV,
— Lf
AY
So
ees
N D
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r\ re,
YW {
/) {/
J
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a
Ss
330
pm, cylindraceae, rectae vel curvatae, biseriatae, in extremitatibus ro-
tundae. Paraphyses filiformes, ramosae, septatae, usque ad 1 pm latae,
ascos superantes ad 8 pm. Excipulum ectalum ex textura oblita, gelatino-
sum, cellulae hypharum pachydermae vitreaeque, usque ad 32 x 5.5 ym. Ex-
cipulum medullatum ex textura intricata, hyphae hyalinae leptodermeae
usque ad 2 pm latae. - Holotypus: In ligno decorticato, Darjeeling (W.
B.), Auge 20, 1980; Raghunandan Sharma, PAN 17406. In herbario universi-
tatis Panjab Cryptogamarum, Chandigarh.
Apothecia scattered, small, saucer shaped, hairy, margin strongly in-
curved, up to 1 mm diam, external surface black, the margin shining due
to the granularly roughened apices of the hairs, hymenium light brown.
Hairs cylindrical, tapering towards the apices, dark brown, granularly
roughened, thick-walled, multiseptate, up to 14-septate and 130 ym long,
cells “up “to (11.5. x 355, pm, wallof céll ups to) 2eum penickaenatiscmocrer
cemented together to form teeth, particularly at the margin. Asci J-,
8-spored, 90-108 x 7.5-9 pm, apex hemispherical, up to 2 pm thick. Asco-
spores hyaline, up to 7-septate at maturity, 15.5-21 x 2.5-3.5 pm, cylin-—
drical, curved, with round ends, arranged more or less biseraitely. Para-
physes filiform, hyaline, branched, septate, up to 1 pm wide at the api-
ces, not swollen, projecting up to 8 pm beyond the ascus tips. Excipulum
differentiated into two zones: ectal excipulum of textura oblita, up to
84 pm thick, cells very thick-walled, up to 32 x 5.5 pm; medullary exci-
pulum of textura intricata, up to 29 pm thick, hyphae hyaline, up to 2.0
pm wide.
ETYMOLOGY: In honour of S. E. Carpenter, monographer of the genus.
HABITAT: On decorticated angiosperm wood.
TYPE\ LOCALITY: India: 6th mile, Darjeeling (W. B.).
HOLOTYPE: PAN 17406, R. Sharma, August 20, 1980; ISOTYPE: CUP-IN 608.
NOTES: Though in some respects this species recalls Xylogramma Wallr.,
its affinities are surely in Crocicreas, though it is clearly distinct
from any of the 54 taxa accepted by Carpenter (1981).
ACKNOWLEDGEMENTS
The senior author is thankful to Dr. W. R. Arendholz, Fachbereich Biolo-
gie, Universitat Kaiserslautern, Kaiserslautern, West Germany for help
with the Latin diagnoses. Financial assistance from the Department of
Science and Technology (DST), India, is gratefully acknowledged.
REFERENCE CITED
CARPENTER, S. E. 1981. Monograph of Crocicreas (Ascomycetes, Heloti-
ales, Leotiaceae). Mem. New York Bot. Gard. 33: 1-290.
MYCOTAXON
VOMREOXV IS NO. 17) opp.) Ooi 54 October-December 1982
a a ee eS
REVUE DES LIVRES
par
G.L. HENNEBERT
Book Review Editor, Croix du Sud 3, B-1348 Louvain-la-Neuve
Belgique
PREDOMINANTLY HOLOCARPIC AND EUCARPIC SIMPLE BIFLAGELLATE PHYCO-
MYCETES, by John S. KARLING, 252 p., 64 plates, Din, hard cover,
1981, J. Cramer, FL-9490 Vaduz, Lichtenstein.
This large size Cramer's edition is appropriate for the publica-
tion of the 64 ful-page plates of line drawings beautifully prepared by
the author for the illustration of the 150 species described and which
so much remind the high quality Thaxter's line drawings.
The species are classified in 7 families on the basis of thallus
structure, developmental cycles and the methods of asexual and sexual
reproduction. As the way of reproduction and of production of resting
spores are unknown in many species, the limits of the genera are not
sharply defined and their classification in coherent families is only
tentative. Such classification is nowaday however provided as an aid
in identification and a means of reference. The distinction between holo-
carpic and eucarpic organisms is not a clear cut and intermediate repro-
duction is found. The expression "predominantly holocarpic" is thus
prefered. Two new species and one newcombination are established. The
31 unnamed species however described demonstrate the state of constant
progress in that field of taxonomy.
THE AGARIC GENERA LENTINUS, PANUS, AND PLEUROTUS, with particular
reference to Malaysian species, by E.J.H. CORNER, in Beihefte zur
Nova Hedwigia, part 69, 169 p., 2 pl., 40° fiel, 8°, hard cover,
1981. J. Cramer, FL-9490 Vaduz, Lichtenstein.
Lentinus Fr. is redefined with skeleto-binding hyphae and relates
with Polyporus sensu stricto. Panus Fr. is defined with long intercalary
or terminal skeletal hyphae without binding hyphae. Pleurotus Fr. is
either dimitic with tapering terminal skeletal hyphae without lateral
branched binding hyphae and with thickened generative hyphae, or monomitic.
Panus and Pleurotus relate with other polyporoid genera. Ten Lentinus
species and one variety are described, 7 of which are Malaysian. Twenty
species and 7 varieties are described in Panus, amongst them 12 species
and the 7 varieties are Malaysian. Pleurotus includes 10 non Malaysian
species and 23 species and 8 varieties from Malaysia. Many of these Ma-
laysian taxa are new.
HIGHER TAXA OF BASIDIOMYCETES, by Walter JULICH, in Bibliotheca
Mycologica nm 65,485 ps; 20 pls. 34 figs, 8°, hardicover,, 1981.
J. Cramer, FL 9490 Vaduz, Lichtenstein. Price: DM 120./150.-
The author proposes a new classification of the Basidiomycetes
based on microscopical characters such as basidia, basidiospores and
S52
hyphal structure. He also emphasizes the phylogenetic relationships be-
tween members of traditional groups like Agaricales, Aphyllophorales
and Gastermycetes in the Homobasidiomycetes. Interesting relationships
are shown also in the Heterobasidiomycetes. In the Gasteromycetes,micro-
scopical studies should also reveal more evidences of affinities. A
general phylogenetic scheme is proposed in which the Cantharellales and
the Auriculariales take the oldest position.
The class Homobasidiomycetes is subdivided in 12 unnamed groups
of orders which are 49 all together. The Tricholomatales, the Entoloma-
tales, the Amanitales and the Pluteales are segregated in the group 6
from the Agaricales (group 7). The Sclerodermatales, the Melanogastrales,
the Leucogastrales and the Tulostomatales are with the Agaricales in
8roup 7. The Russulales are grouped with the Bondarzewiales and the
Hericiales in group 3. The author's hope is that the resulting classifi-
cation 1S a more natural one.
In the special part of the book, characteristics and affinities
of each family and higher taxon, many of which are new, are detailed and
illustrated with line drawings and SEM spore photographs.
THE RESUPINATEN PHELLINUS-ARTEN IN MITTELEUROPA, mit Hinweisen auf
die resupinaten Inonotus-Arten und Poria expansa (Desm.) (=Polypo-
rus megaloporus Pers.), by H. JAHN, Bibliotheca Mycologica n° 81,
152, p.., 21 fie., 61 phot.,’8 3 paperback, reprint {98le J. Ceamere
FL 9490 Vaduz, Lichtenstein.
The text consists of a paper reprinted from the Westfalische Pilz-
briefe, 4(3-6):37-108, 1966-1967 followed by an original supplement,
subtitled "Nachtrage 1967-1981 (Fig. 13-21)" of 43 pages. In these papers
the author demonstrates the anatomical differences between the two rela-
ted genera Phellinus and Innonotus. He provides comments on the charac~
teristics and the ecology of 16 Phellinus species and 5 Inonotus species.
Keys are proposed for both genera. Boletus expansum Desm. ( syn. Polypo-
rus megaloporus Pers.) was renamed Poria expansa in 1967 and later
Donkiopora expansa by Kotlaba and Pouzar; this species is also considered
for its similarities to Phellinus.
STRUKTUR UND FUNKTION MITOCHONDRIALER DNA BEI PILZEN, by Hels KUCK,
in Bibliotheca Mycologica n° 84, 148 p., 19 fig., 8°, paper back,
1931... J. Gramer, EL=9490,,. Lichtenstein.
In this thesis, the author reports the isolation and characteriza-
tion of the mitochondrial DNA from two fungi, a yeast, Saccharomycopsis
lipolytica, and a filamentous ascomycete, Podospora anserina. In both
species the mitochondrial DNA is a circular molecule with a GC contents
(27.5 and 34.0 Z% respectively) lower than that of the nuclear DNA. The
length of the molecule reaches 15.4 um in Saccharomycopsis lipolytica
and 32.8 um in Podospora anserina. In the later species, the mtDNA is
only recognized in a juvenile state but changes into a plasmid-like DNA
of maximum circular length of 9.6 Lm composed of monomere of 0.75 um in
length in the senescent state of growth. The author has been able to
clone such pl1DNA in Escherichia coli.
THE BIOLOGY AND CULTIVATION OF EDIBLE MUSHROOMS, by S.T. CHANG and
W.A. HAEYES, xxii + 819 p.,8°, hard cover, 1978. Academic Press,
New York.
Re)
Since 1965, at the International Mushroom Congress in Amsterdam,
the cultivation of mushrooms other than Agaricus bisporus started in
Western Europe, while several species where already cultivated in the
Eastern world, Japan and China, f.i. Tricholoma matsutake.
This book is the first comprehensive treatise available on the
general biology and the culture technics of the different kinds of
mushrooms now in cultivation. These are the fungi that grow on almost
fresh plant residues (Lentinus, Pleurotus, Flammulina, Auricularia,
Pholiota, Tremella, Agrocybe, Ganoderma, Coprinus), the fungi that grow
on only little composted material (Volvaria, Stropharia, Coprinus), the
fungi that grow on very well composted material (Agaricus), the fungi
that grow on soil and humus (Lepiota, Lepista, Morchella, Gyromitra) and
the mycorrhizal fungi (Boletus, Cantharellus, Amanita, Tuber, Morchella,
Lactarius, matsutake). The book is general and specific. It bridges the
Sap between researchers and growers. It shows the problems and the
progress in all aspects. Written by 32 contributors from eleven countries
it deals with many different ways of mushroom cultivation. Nutritional
value, medical effects and economical aspects of the increasing consump-
tion of mushrooms are also considered.
MONOGRAPH OF THE PYTHIUM, by A.J. VAN DER PLAATS-NITERINK, in
Studies in Mycology n 21, 242 p., 103 fig., 8°, paper back, 1981.
Centraalbureau voor Schimmelcultures, Baarn, The Netherlands.
HELO.
This revision of Pythium Pringsheim is mainly based on living cul-
tures preserved at the C.B.S. and a critical review of 1133 publications.
85 species are recognized and described alphabetically, 64 from living
strains. In addition, two highly cellulolytic species are treated in an
appendix. The species P. buismaniae and P. macrosporon are new. 65 remai-
ning species are classified as uncompletely known, doubtful or to be
excluded. The 87 accepted species and 13 uncompletely known species are
keyed out dichotomously. Off those, 81 are homothallic, 9 are heterothal-
lic, 1 species and 5 unnamed group-species have no oogonia but sporangia
and 9 species have neither oogonia nor sporangia. In such a large and™
difficult genus, a synoptic key might have been helpful.
FUNGAL PHYSIOLOGY, by David H. GRIFFIN, xiv + 383 p., ill., 8°,
hard cover, 1981. John Wiley & Sons, Wiley-Interscience, 605 Third
Avenue, New York NY 10158.
The author has made a very needed synthesis of present knowledges
on the physiology of the fungi. In such a field of steady progress, he
provides us with a critical analysis of the different experimental ap-
proaches used by researchers. After chapters on the specific biochemistry
and molecular architecture of the fungus cell, the author considers the
primary and secondary metabolism. Chapter 5 to 8 deal with the growth,
its rate and regulation, its chemical requirements, the transport and
absorption of the nutrients and the response to the environmental factors.
Chapter 9 to 12 deal with the reproduction: the effects of environmental
factors on the spore formation (including circadian rythms), the bioche-
mical and genetical regulation of the spore formation, the dormancy and
germination of spores and the interesting aspects of syngamy, like the
hormonal regulation in sexual interactions. Final chapters describe the
behaviour of the fungi in natural attacks and provide a ratioral basis
for the choice of fungicides. The text is precise and well documented
by data and graphs from the literature. The author also points out the
gaps in the present knowledge from a dynamic point of view.
334
CHAMPIGNONS DE SUISSE, Contribution a la connaissance de la flore
fongique de suisse, TOME 1, LES ASCOMYCETES, Photographies en
couleurs, descriptions et dessins d'observations microscopiques de
390 espéces de Suisse centrale et particuliérement du canton de
Lucerne, by J. BREITENBACH and F. KRANZLIN, French translation by
J. .KELLER, 310 p., 390 fig., 390 col. “phot. 74 9 hararcover loci
Ed. Mykologia, CH-6000 Lucerne, Switzerland. SFr 118.-
This album is a modern and scientific account of the Ascomycetes
from Central Switzerland, but common to all European countries. It is
the first volume of a series prepared by the authors with the collabo-
ration of members of the Société Mycologique de Lucerne. It results
of a methodical and precise work procedure in collecting, describing,
illustrating and preserving the fungi which will surely satisfy the
scientist and traine the students. 384 Ascomycetes species are treated
in a taxonomical order, within 37 families. Named by their Latin and
French names, each species is-described and illustrated from an accura-
tely indicated collection which is deposited in herbarium. Macroscopical
features are shown on a high quality color macrophotograph and microsco-
pical structures exhibited in line drawings. The habitat of the species,
taxonomical comments and pertinent literature are also given. The keys
are di- or multichotomous and require the use of a microscope.
That book is a joy for the reader also for its design and color
printing. It is an invaluable contribution towards the diffusion of a
scientific knowledge of these fungi.
COMPENDIUM OF SOYBEAN DISEASES, by James B. SINCLAIR, Ed. 2d ed.,
104 p., ill., 4°, paper back, 1982, The American Phytopathological
Society, 3340 Pilot Knob ‘Road, St» Paul s.Minnesota 5512) 7 Salto
This second edition, revised with the aid of 76 world authorities
in the field, has new sections including one on diseases caused by myco-
plasmalike organisms, one on diseases of unknown origin and one on di-
S€ase control strategies. The other sections have gained from more in-
formations, references and illustrations. The 44 fungal diseases of
soybean take a major importance in this Compendium.
AN ANNOTATED CHECK-LIST OF FUNGI CAUSING POSTHARVEST DISEASES OF
FRUTS AND VEGETABLES IN ISRAEL, by Rivka BARKAI-GOLAN, Special
Publication n°194, 36 p., paper back, 1981. Division of Scientific
Publications, The Volcani Center, Bet Dagan, Israel.
This little paper proyides with an alphabetical listing of the
concerned fungi with indication of their host, the kind of symptoms, a
brief account on the distribution and the importance of the decay.
MYCOTAXON
VOR le NO. I pps 335-3359 October-December 1982
Ie a ee oe a ee
NP OeT 7 C#e
INTERNATIONAL MYCOLOGICAL ASSOCIATION
RECORD OF THE BUSINESS MEETING HELD ON 26 AUGUST 1981
DURING THE XI11 INTERNATIONAL BOTANICAL CONGRESS, SYDNEY
Agenda
1. President's remarks
2. Apologies for absence
3. Secretary's report
4 Treasurer's report
5 Committee reports
(a) Nomenclature Secretariat
(b) Asian Mycology
(c) Latin American Mycology
6 IMA Statutes
7. Third International Mycological Congress
8. Structure of IUBS
9. Relationship with IUMS
10. XXI General Assembly of IUBS
Hee atavSon., Gr ice
12. Any other business
1. President's remarks
1. Professor C.V. Subramanian (President, IMA), welcomed members to the
Business Meeting.
2. He first paid tribute to three outstanding mycologists who had
passed away since IMC2, Professor Ralph Emerson, Professor F.K. Sparrow,
and Dr Luella K. Weresub. Two minutes silence was observed by the whole
Meeting.
3. Professor Subramanian welcomed the opportunity of mycologists parti-
cipating in the Sydney Congress to meet together and the possibility of
reporting on recent activities of the IMA to a wider audience than was
usually possible.
4. He then proposed that the Agenda for the meeting, which had
previously been circulated to affiliated organizations and members of the
Executive Committee, be approved, and this was adopted by the Meeting.
2. Apologies for absence
5. Apologies for absence were received from three members of the Exe-
cutive Committee, Dr J.A. von Arx (Treasurer, IMA), Professor J. Webster,
and Dr G.C.A. van der Westhuizen.
336
3. Secretary's Report
6. Dr D.L. Hawksworth (Secretary, IMA) reported that since the IMA
officers and Executive were elected at IMC2 in 1977, Professor R.A.
Emerson (USA) passed away, and was replaced on the Executive by
Professor A. Skirgiello (Poland). There have been a number of changes
of addresses and a current list of the members of the Executive, which
will serve until IMC3 in 1983,was tabled.
7. %In 1978 the Executive considered submissions from the Mycological
Society of Japan and the Mycological Society of India to host IMC3 and
agreed that Tokyo was the most suitable centre. I would like to record
the gratitude of the IMA to both Societies for their invitations. 1983
was selected as a date so that there would be a full year between the
present Congress and IMC3. Precise dates were fixed to avoid a direct
clash with the ISPP Congress in Australia the same year.
8. Copies of the Resolutions from IMC2 were circulated to national
and international bodies, published in the IUBS Newsletter, and
endorsed by the XX General Assembly of IUBS in Helsinki in 1979.
9. Professor Subramanian and Dr Hawksworth attended the XX General Assemb]
of the International Union of Biological Sciences (IUBS) “to present a
case on behalf of the Executive Committee for the recognition of an
increased status for mycology within IUBS. As a result of discussions
held in Helsinki, it was agreed to re-name the Division of Botany as
the Division of Botany and Mycology, and further to create a Section
of General Mycology, based on IMA, to replace the former Commission on
Mycology. Professor Subramanian was also elected to the IUBS Executive
as one of the representatives of the new Division at the Congress, “The
changes made at Helsinki! pave the road for a strengthening of the voice
of mycology in world biology.
10. Since IMC2, the Australian Plant Pathology Society, the Inter-
national Association for Lichenology, the Society for Human and Animal
Mycology, and the Societas Mycologica Fennica have all become affiliated
to the IMA. A mutual affiliation With the International Association.
for Plant Pathology has also been agreed in principle.
4. Treasurer's Report
11. Dr J.A. von Arx (Treasurer, IMA) had provided a statement of the
current financial position for the information of the Meeting which
Showedrancunrenteoalance! Of shill 27 s572 vat tad ainyehO Sle
5. Committee Reports
(a) Nomenclature Secretariat
12. Professor R.P. Korf (Member, Nomenclatural Secretariat), read a
Report of the activities of the Secretariat to the Meeting.
13. The formal proposals for improvement of the treatment of fungi in
the International Code of Botanical Nomenclature prepared by the IMA
Nomenclature Secretariat, following reports of its Committees and de-
bates at IMC2, were published by the Chairman of the Committee, Dr K.T.
van Warmelo inTaxon 28: 424-431 (1978).
53/
14. These proposals were considered by the Nomenclature Sessions prior
to the Congress. It was unanimously accepted that the starting point
date for all fungi would be changed from 1801 or 1821 to 1753 (with
special provisions for names in the formerly cited "Starting-point
books"). The completely revised version of the Rules governing the
names of fungi with pleomorphic life cycles was also accepted. Pro-
posals to introduce living types into the Code failed, but one of the
IMA's Recommendations was adopted and the Session recognized the problem.
There was an impression that fresh proposals would be viewed sympath-
etically.
15. It had further been agreed in the Nomenclature Sessions of the
Congress that three posts in the Special Committee for Fungi should be
left vacant until IMC3. At IMC3 it may be proposed that this enlarged
Special Committee should act as the Nomenclature Secretariat and appoint
committees as required. Such a move might well eliminate the need for
the present dual structure for considering problems of mycological
nomenclature.
16. With regard to the question of the acceptance of living cultures
as types, the present Meeting recommended that the Special Committee
and Nomenclature Secretariat should cooperate with algologists in form-
ulating fresh proposals.
(b) Asian Mycology
17. Professor Subramanian reported that he hoped to arrange a dis-
cussion meeting for Asian mycologists during 1982.
(c) Latin American Mycology
18. No report of recent activities was received.
Gis IMA Statutes
19. The Meeting considered the draft Statutes of the IMA which had
been drawn up following IMCl1 in 1971, but had not been formally adopted
by a subsequent International Mycological Congress.
20. The present structure had meant information too rarely percolated
to individual mycologists, and there was a strong feeling, especially
amongst Australian participants, that some form of individual member-
ship should be retained even in countries that had national affiliated
organizations.
21. The Executive Committee was instructed to take note of the views
expressed and the Secretary agreed to prepare a revised set of Statutes
for their consideration. Participants undertook to send any further
comment they wished to be considered to the Secretary by 1 November 1981.
Te Third International Mycological Congress
22. The Meeting welcomed Professor K. Tubaki (Secretary-General, IMC3),
who outlined the arrangements being made by the Japanese Organizing
Committee. Professor N. Hiratsuka had been elected President, and
Dr K. Iwata Vice-President of the Committee. The congress would be
based on the Keio Plaza Hotel (Shin-juku, Tokyo) but it was expected
558
that most participants would stay in an adjacent business-class
hotel. The First Circular had now been printed and copies were
distributed to the Meeting. The Circular included a list of major
topics to be included in the programme. A registration fee of about
US $ 150 was anticipated. The Organizing Committee looked forward
to the visit of the Secretary of the IMA to Japan following the
Sydney Congress.
23. Professor T. Ahti (President, International Association for
Lichenology), hoped that a lichenologist would be included in the
Organizing Committee.
See SeLuceune. Of UBS
24. Professor K. Esser outlined the discussions on the structure of
IUBS that had taken place since the XX General Assembly of IUBS
in 1979 (see para 9 above). The division of Microbiology had now
left IUBS and had been formed into an International Union of Micro-
biological Societies (IUMS). The board of the Division of Botany
and Mycology of IUBS had also considered structure at a meeting on
23 August during the Sydney Congress. The Board resolved to urge
IUBS to maintain the present Division of Botany and Mycology, to
recognize the Chairman of that Division as the voting member of the
IUBS Executive, and to delegate budgetary responsibility for the
Division on the Board. It was further proposed that the President
of the Division would be that of each individual International
Botanical Congress and that he would serve for three years before to
three years after the Congress.
9. Relationship with IUMS
25. The Meeting noted the establishment of IUMS, and that it included
a Division of Mycology. The Secretary summarized a letter he had
received from Dr. Iwata (Chairman of that Division) hoping that the
two organisations would collaborate in fostering actual co-operation
in developing mycological research in all its fields. After some
discussion it was agreed that the informal links that had previously
existed when the Division was a Section within the Division of Micro-
biology of IUBS should be maintained, but it was not considered to
be appropriate to enter into any formal relationship with IUMS at
that time.
10. xII General Assembly of IUBS
26. No proposals had been received, but the Meeting agreed that the
IMA should support any applications received from the International
Association for Lichenology (IAL) or the International Society for
Human and Animal Mycology (ISHAM) for recognition as Sections with the
IUBS Division of Botany and Mycology.
27. It was agreed that the President, Secretary and (if possible)
Dr.S.J. Hughes Vice-President IMA should represent the IMA (section
for General Mycology) at the General Assembly.
Le Re,
Me eLtalson OLfice
28. The secretary regretted that further progress had not been made
in efforts to seek funds to establish a Liaison Office for world
mycology as recommended by IMC2.
12. Any other business
Professor K. Esser informed the Meeting that the XIV International
Botanical Congress would be held in Berlin in 1987. The venue
would be a conference centre which could contain 7000 delegates and
Professor Esser invited the IMA to hold IMC4 conjointly with that
Congress. After some discussion, the Meeting agreed that Professor
Esser should explore the possibilities further. This, and any other
offer to host IMC4, would have to be considered by the Executive
Committee of IMA prior to IMC3. The Executive would hope to be able
to make a firm recommendation to the IMA General Assembly to be held
during IMC3.
30. In closing the Meeting, the President thanked all the mycologists
who had made time in their busy Congress Schedules to participate,
and also paid tribute to the support he had received from the Officers
of the IMA.
Wye) jweneatlk WSksyZ Dr. D.L. Hawksworth
Commonwealth Agricultural Bureaux,
Farnham Royal,
Slough SL2 3BN.
Uke
Secretary, IMA.
MYCOTAXON
VOL. UAV TeiNo fae pe 540 October-December 1982
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TAAGA, MY. 14853
Volume. XVI January-March. 1983 No.
MYCOTAXON
AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION
OF RESEARCH, ON. TAXONOMY & NOMENCLATURE OF FUNGI) & LICHENS
Z
CONTENTS
Sanctioned epithets, sanctioned names, and cardinal principles
Ii ers ol ana eb AMM tat MOMS Vik eeh a webracol blabcl alk RICHARD (Po KORF
On the genera Cochitobolus and Pseudocochliobolus..... J. LU. ALCORN
PEWS Pe eS OT SR AE LI ies Konrauieh UCL RPL gone Qi atlnr dieoids G1 col dd was Nw BENYAMINI
A new Japanese species of Neocosmospora from marine) sludges,
SEIICHT UEDA ‘AND .SHUN-TCHT UDAGAWA
New records of hypogeous Ascomycetes in Arizona... JACKS.) STATES
Studies in the genus, Phoma. 1. Phoma amertecana sp.nov.
G. MORGAN-JGNES AND JAMES) \F.. WHITE
Ay ROW. SROCLES Of Amo t Pai. Wis cieidu says bite AVR Aa asta Woes DAVID: TL. JENKINS
Descriptions of new species and combinations in: Mécrosphaera
SUG MOTE) 8 aR IANS ERG: ALN SAR AAA FL) MMA RO BREN nD SU PH UWE BRAUN
Taxonomic notes on some/powdery mildews) (11)... vel ee etn UWE BRAUN
Gigaspora reticulata: a néwly described, endomycorrhizal fungus
from New England.
R, E. KOSKE, DIANE DOUD) MILLER ‘AND CHRISTOPHER WALKER
A new thermophilic species of Myceltophthora.
TAKEYOSH! AWAO AND SHUN-ICHT. UDAGAWA
Operculate Discomycetes’ from Rana (Norway) 5. Rhodoscypha gen.
nov. and Rkhodotarzetta gen. nov.
HENRY DISSING!) AND STGMUND SIVERTSEN
Basidiomycétes Aphyllophoralés epitheloides étalés.
J.) BOIDIN AND. P. LANQUETIN
New South American resupinate polypores. 2.05... MARIO, RAJCHENBERG
Colletotrichum gloeosportotdes (Penzig) Penzig et Saccardo.
S.. Rs. PENNYCOOK
Ectomycorrhizae of selected conifers growing in'sites which
support dense growth of bracken fern.
JAN -ACSAT AND DAVID L. LARGENT
Mycorrhizae of Arbutus menetestz Pursh. and Arctostaphylos
manzantta, Parry in northern California.
JAN ACSAT AND DAVID, Lis LARGENT
Check-list of Romanian Peronosporales.
O. CONSTANTINESCU AND G3' NEGREAN
[CONTENTS continued overleaf]
54)
SOD
380
Por
396
AQ3
4i4
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A25
429
436
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ISSN 0093-4666 MYXNAE? 2642), 3412-582 (1983)
Library of Congress Catalogue Card Number 74-7903
Published quarterly by) MYCOTAXON, Ltd., P.O; Box 264, Tthaca NY 14850
For subscription details, availability in microfilm and microfiche,
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CONTENTS (‘continued
Sarctnulelia bankstde gen) et sp. nov.’, a Coelomycete with a
uniague method of conidial dispersal.
Bec SUTTON AND J. LG. ALCORN 557
Cultural studies in Kypochntetum (Corticiaceae, Basidiomycetes).
NILS HALLENBERG 565
Notice: MSA plans) for MC TE CCAS SPN GTS Ae SPU LN BE TORE Gy OE EE MURTY A pte ep 572
AUPE O TY VENETO e Gay ikl ORC ani Ra ats ht ce We Oye: Whe MoU RS RO MN eeradiotany Strata bad oe st 57S
ENDED C0) Mang OUS. Sitd) CE CRem NEE Ae CoML Rs Lie eA URL HAR Ageia 576
BRAC Sar ye Brahe able amte ERR Sl fe cuted ok aiietidy WT Gibbet bi wr oF AERIS pa iO OL aes Ma aU Se ae et a 582
MYCOPAXON publication dates J) bs sama 1G (hoa nett ea Oe couhen Sate 582
REN MEWOR SU ec Cente Bate Ue T ay rane cOLEe ae cian Milas 1s nA RA Seal vt
IMYCOTAXON for Cetober-December’ 1982) (16: 1~340)
was issued October 27, 1982]
Fag ES a
MYCOTAXON
Vobwenvil No, 2. pp. 341-352 January-March 1983
SANCTIONED EPITHETS, SANCTIONED NAMES, AND
CARDINAL PRINCIPLES IN ": PERS." AND ": FR." CITATIONS
RICHARD P. KORF
Plant Pathology Herbarium, Cornell University, Ithaca, NY 14853 USA
ABSTRACT
A historical perspective on the International Code of
Botanical Nomenclature is given on (1) the establishment
in 1910 of later starting-point works for fungal nomencla-
ture (Persoon's Synopsis Methodica Fungorum of 1801 and
Fries's Systema Mycologicum of L82t-1832)5 “and; .(2) the
ditemma of 1950-1981 in citing author citations for early
species names once a 1 Jan. 1821 starting-point date had
been established for ‘fungi caeteri.' The concept of sanc-
tioned names (and epithets), introduced into the Code at
the Sydney congress in 1981, is explained. An urgent plea
is made to mycologists to find holotypes or to designate
lectotypes or neotypes of sanctioned names so that taxono-
mic mycology can proceed on a sound footing. Some princi-
plese an, sehe ypractical Japplication of jthe "“: Pers." and
":; Fr." system of notation for sanctioned names and epi-
thets, also introduced at Sydney, are elucidated.
BP VSRIER HISTORY OF LATER STARTING-POINT BOOKS AND DATES, AND
OF THE ORIGIN OF "SANCTIONED" NAMES
At the 1910 (Brussels) International Botanical Congress
(IBC) various later starting points for the nomenclature of
plants were adopted where use of Linnaeus's Species Planta-
rum was felt ill-advised. For fungi four different starting-
point works were named: Linnaeus's Species Plantarum (1753)
was accepted for a starting- point book for myxomycetes and
for lichens; Persoon's (1801) Synopsis Methodica Fungorum
for rusts, smuts, and gasteromycetes; Sternberg's (1820)
pora ;aer |Vorweldt for ali fossil plants’ (including fossil
fungi); and Fries's 3-volume (1821-1832) Systema Mycologicum
Por-allsother fungi (termed the ''fungi caeteri").
All fungal groups, except fungi caeteri, had a single
volume as the starting-point book. Between 1910 and 1950 an
meten lively debate arose over how to interpret the 1821-
3032 date-spread as a starting point for fungi caeteri. A
few authors held that the starting point should be 1821,
and that all names published after that date were validly
published. Others held that names published prior to 1832
had no status and were not validly published except in the
Systema itself. Still others assumed each group of fungi
Mao its ‘starting point on the date that Fries took up that
342
group in the Systema. Thus agarics and aphyllophoralean
fungi would mostly date from 1821, as would the Geogloss-
aceae of the Discomycetes; many other Discomycetes would
date from vol. 2(1) of the Systema in 1822; the remaining
Diseconycetesm@en2tZy in 1623s. ete,
Because precision in dating nomenclatural matters is
essential, a major change was enacted at the 1950 (Stock-
holm) IBC in which ‘actual or arbitrary publication dates
were set for starting-point works, and nomenclature for
each of the groups was now defined as beginning on a
particular date (rather than book). The dates which concern
us were 1 May 1753 for ,Linnaeus's (Species Plantarumes.
Dec. 1801 for Persoon's. Synopsis, 31 Dec.” 1820) formes vera
berg rs Flora; and, 1° Jan. 1821) for “volume; ore@ureuue
Systema. (The other volumes of the Systema were not arbitra-
rily dated.)
The whole purpose of adopting later starting points
had been to attain stability in nomenclature, recognizing
that, in the case of fungi, there were two early authors
Persoon and Fries, who had brought some semblance of order
out, of) the chaos’ of their pasts. “The intent in 2910gwaceue
build upon their taxonomy (and thus their interpretations
of the taxonomy and nomenclature of earlier works). With
the adoption of a starting-point date of | 1) Jans toZ20 pea
those fungi caeteri in volumes 2 and 3 of the Systema were
deprived of a starting-point book for reference. Since this
was clearly ‘at’ odds with ‘the intent) of > having spr ic ass
Systema serve as a later starting point, the 1950 IBC
adopted a method of providing priviledged status to the
names in those two volumes. They extended that priviledged
status also to the names used by Fries in the 2-volume
Elenchus Fungorum (1828), correctly seen as a true supple-
ment to the Systema. The exact wording adopted for Art. 23f
of the Stockholm Code was:
f. FUNGI CAETERI, 1 Jan. 1821 (Fries, Systema Mycologicum
Vol. 1). Vol. 1 of the. Systema is treated :as ‘having ap-
peared on 1 Jan. 1821, and the Elenchus Fungorum (1828) is
treated as part of the Systema. Names of FUNGI CAETERI
published in other works between the dates of the first
CVol.- 1), and last -(Vol. 3 part 2 and “ndex )Epartceoneene
Systema which are synonyms or homonyms ! of names of any of
the FUNGI CAETERI included in the Systema do not affect
the nomenclatural status of names used by Fries in this
work.
The: wording was not. felicitous) since theremppewane
definition of what the “nomenclatural, status" of (Fries
mames is. The French text is perhaps clearer "osgeneaae
point de répercussion sur la nomenclature des noms utilisés
par Fries dans son ouvrage.'' The names in vols. 2 and 3
(and in the Elenchus) were thus accorded a special kind
of priority as protected or priviledged, names, and) tue
A reminder to the reader: a synonym is a different name for the same taxon based on the
same or on a different type; a homonym is the same name based on a different type.
343
whole Systema was in some ways still a "starting-point
book.'' For these priviledged names Donk (1961) coined the
term "sanctioned" names. He also suggested adding an indica-
Prom 01. such Sanctioned .status,'':-Fr.:'", to -those. names
which were validly published prior to the date that they
were streated by Fries in the Systema: |Since: valid publica-
BronyoL fungi cacteri “did not) begin until 1 Jan. 1821, the
": Fr.'' notation was only to be applied to names published
in other works between 1821 and 1832 (index) that were
adopted (in some form) by Fries in the Systema/Elenchus. ]
Though Donk's formal proposal for this system of notation
was rejected at the next IBC, many workers, myself included,
found it so useful that they often adopted this system in
their publications.
THE 1950-1981 DILEMMA: AUTHOR CITATIONS
Witmeeene adoption in 1950 of 1 Jan. 1821 as the start-
ing-point date for fungi caeteri, new problems in citing
authorities arose. A single example is chosen here to illus-
trate how the many variations in permissible author cita-
tions came into being.
beziza. araneosa Bull’ 1791, Gf it» had been ‘a, plant
with a 1753 starting point, would have complied with all
Beesevequirements “for “valid publication. Since it is a
member "or the fungi caeteri, it was "devalidated" (a term
eorned | by Donk, 1957)! by provisions of later ‘starting
Ponts. (Fries accepted this species in Syst. Myc. 2(1): 69,
Beee-eerrior to 1959 the International Code of Botanical
pemenclature (ICBN) permitted citation of this name in
three ways:
(i )—Pezi za araneosa- (Bull. | Fr.
(2) Peziza araneosa Bull. ex Fr.
(3) Peziza araneosa Fr.
Sie "tirst “two options cléarly identified the source of the
name, and thus of its type specimen; the third obscured the
origin of the name. The foolishness of having two different
ways of indicating a pre-starting-point author, square brack-
ets and ''ex,"’ and the problem caused by thus introducing a
) different meaning for ''ex'' than used elsewhere in the ICBN,
has been detailed elsewhere (Donk, 1957). In the 1959 and
#764 Codes only citations (1) and (3) are approved. A
Beversal of this occurred in the 1969 and 1975 Codes, where
only options (2) and (3) are approved. [The change was made
(unwisely!) by the Editorial Committee, and not voted upon
by the Congress. A proposal by Compére (1973) to restore
Bquare brackets instead of "ex'’ was not accepted at the
Beningrad (1975) IBC..] ary
Witt tne adaption: ‘in’ 1990) at) Stockholm “of “a” start—
goe-point date of 1 Jan. 1821 a wholly new problem arose:
whosoever first adopted the name after that date became the
Walidating author (or, better, in Donk's terminology, the
"revalidating" author of a devalidated name). The immense
Mmeitficulty of ascertaining the: name of this validating
344
author was not given serious consideration by those who
adopted the 1950 ICBN. The problem has been discussed at
length elsewhere (Korf and Kohn, 1979, 1980; Demoulin et
al. 1981) and need not be restated here. Under the 1950
Code Bulliard's name (above) had special status since it
was ‘sarnctioned™ by its use in Fries’ s Systema. Buewsene
author citation was no longer that of the pre-1950 Codes,
for Fries ‘only took’ up the name in Late 1822, "and at sleace
four authors used it in 1821 (and at least one ‘others aucunen
earlier in 1822 than Fries).
Extensive and painstaking bibliographic research over
the years by such experts as Donald P. Rogers and Ronald H.
Petersen (see Korf, 1982a) has led to partially complete
sequencing of the various books published in 1821 (and, to
a much lesser extent, those published in 1822 and later).
But for some 1821 books and articles an exact date of publi-
cation (and therefore their position in the sequence) will
never be able to be established. (One could, of course,
legislate arbitrary dates of publication for all of then ge
Of the four known competing treatments in 1821 that adopted
Peziza araneosa Bull., we know (Petersen, 19/5) that St.-
Amans's Flore Agenaise was published between the 20th and
28th of April, Mérat's Nouvelle Flore between the 9th and
15th of June, and Purton's Appendix after August Ist28ue
Nocca and Balbis's Flora Ticinensis is, in,Petersen’ saworace
"unsequencable."
The 1950-1975 Codes left us with four’ possibly correct
eHEaclronus ;
(4) Peziza araneosa Bull. ex St.-Amans
(5) Peziza araneosa Bull. ex Nocca & Balb.
(6) Peziza araneosa St.-Amans
(7) Peziza araneosa Nocca & Balb.
For none of these was Fries's name to have any part in the
citation, for his action in 1822 was after the jstartinge
point date. Citations (4) and (5) had at least the meriGyom
telling us where the type of the name is to be found
(Bulliard's specimens if extant, publications, figuresie
Citations (6) and (7) make matters quite abstruse, since
neither St.-Amans's nor Nocca and Balbis's works are avail-
able in many libraries, nor were they cited by) Friecs
Mérat, or Purton. On the contrary, even if Bulliard’s ‘wor
is not in many libraries, it is cited by Fries and by every
one of the above. authors,.,as 1t was: ‘the basis (of, ae cies
treatments. (It is, moreover, now easily available on micro-
fiche.)
And yet a further question haunted us: did Rogers and
Petersen really exhaust the 1821 literature? Would someone
unearth another obscure 1821 work using Bulliard's name?
I am intentionally ignoring two additional citations, (4a) Peziza araneosa [Bull.]
St.-Amans and (5a) Peziza araneosa [Bull.] Nocca & Balb., permitted under the 1950
and 1954 Codes and demanded instead of (4) and (5) by the 1959 and 1964 Codes.
345
For example, a publication by Apocryphal, either in Jan.-
Matremio2i= or of undeterminable, date within’ 1821, would
yield two additional possibilities:
(8) Peziza araneosa Bull. ex Apocr.
(9) Peziza araneosa Apocr.
No "stability" in nomenclature arose from these changes in
the Code in 1950: indeed, mycologists who wished to follow
the Code were forced to leave their field work and their
microscopes and to head for the dustiest library shelves to
look at musty local floras, emerging finally with an author
citation that - at best - "might" be the correct one!
THE 1981 CODE: SANCTIONED NAMES AND
A UNIFORM STARTING POINT
In an effort to obviate the drudgery of sterile library
searches that led to sometimes conflicting, sometimes unre-
solvable problems, a subcommittee of the Nomenclature Secre-
tariat established in 1971 by the International Mycological
Association made a series of proposals (after years of work
and good-intentioned if not always good-natured in-
fighting) that resulted in changes in the ICBN enacted at
sydney during ‘the 19381 IBC. In “brief; these restored
Linnaeus's Species Plantarum (1 May 1753) as the starting
Deinvessor all recent’ fungi. (fossils, still start in 13820),
but at the same time gave protected status to the names
adopted by Persoon in the Synopsis (1801) for rusts, smuts,
and gasteromycetes, and to those adopted by Fries in the
Systema/Elenchus (1821-1832) for fungi caeteri (excluding
myxomycetes). Donk's term ''sanctioned'"' was employed for any
name "adopted" in those named works in the respective
eroups:. Donk's proposed "'?"Fr." notation (for indication of
a name ''revalidated" by some other author than Fries that
had, at a later date, been adopted in the Systema) was
taken up in a modified form, since there are now two
sanctioning works. (Persoon's names in the Synopsis were
published under 1950-75 Codes on their own later starting
Porn cate, 31° Dec. 801, and had automatic’ priority; to
maintain that priority, sanctioning status was necessarily
extended to that work.)
What did this do to our example fungus? It changed the
Status of “Bulliard's' 1791 name from “not validly published"
Bo, “validly published," -since it appeared after 1 May 1753.
By examination of the Systema we discover that Fries ''sanc-
tioned" the name. We need never look in the library for the
Botke Oe yot.-Amans, Mérat Purton, or Nocea, and Balbis in
regard to this fungus, for they no longer have any nomen-
elatural impact; their .sole claim to our current interest
lies in distributional records or possible misidentifica-
tions. Likewise, we need never fear the discovery of ''Apocry-
phal's" 1821 treatment. The new Code now permits only two
citations:
(10) Peziza araneosa Bull.
(11) Peziza araneosa Bull. : Fr.
346
For many purposes the first will suffice. For certain situa—
tions the latter will be better, for it conveys an addition.
al bit of information: "this name has been sanctioned by
Fries.'' As we shall now see, sanctioning is a _ two-edged
sword, having to donot: only with “priorability of tnamec,
but also with typification of names.
SANCTIONED NAMES:
PROTECTED AGAINST SYNONYMS AND HOMONYMS
Under .the 1950-1975. Codes, names of . tunel, caeterns
adopted by Fries in vols. 2 and 3 of the Systema and in the
Elenchus were granted a priority over competing names (i.e.,
names at the same rank) published after 1° Jan. 1821 “and
before the index to vol. 3 (1832), whether these were
synonyms or homonyms. Under the 1981 Code, all those names
not- only retain that priority status,” but) now simp
status extending all the way back to 1/753 has been extended
to them and to all of the names of fungi caeteri adopted
(sanctioned) by Fries’ in vol. 1 of ‘the ‘Systema and valsarca
all of the names of rusts, smuts, and gasteromycetes sanc-—
tioned by Persoon in the Synopsis (1801). Sanctioned names
are protected only at the rank assigned them by Persoon or
by Fries in the sanctioning work, since priority is always
rank-limited under ICBN (Art. .60.1.."... In no case does
name or epithet have priority outside its own rank").
TYPIFICATION OF SANCTIONED AND UNSANCTIONED NAMES
A very significant aspect of the changes adopted at
Sydney was a further attempt to attain stability in applica-
tion of sanctioned names. The ICBN is based on the type
method, and though specific and infraspecific names have
long been recognized as having specimens as their types,
the Sydney Code makes amply clear that the names of all
taxa, e.g., generic names, ultimately have specimens which
fix the. application of the name. For many early fungal
names type specimens no longer exist, and for unequivocal
application of a mame, nomenclatural stability requires
us to propose a neotype :to fix the application of sucm
names. Words and drawings cannot be sectioned, nor can
reagents be applied to them to determine reactions. Under
the Sydney Code the type of a sanctioned name may be selec-—
ted not only from the original author's material, but alse
from the material of the sanctioning author or from material
he cites by reference. Sometimes an early author's herbarium
yields one or more specimens bearing the name in question,
but their status as holotype or syntype material may be
open to serious question. It should now be possible dm
most cases to choose a lectotype, or, far more frequently;
a neotype, which corresponds to current usage of a name,
as viewed through the eyes of the sanctioning and the origi-
nal authors. When these two disagree, the choice should be
based on whichever will cause the least discomfort and
disruption of taxonomy. Neotype specimens will be able
to be sectioned, examined, and characterized. A great advan-
tage of these neotypes over the generally "lost" original
materials is that they can be abundant collections, well-
347
dried, and adequately characterized when in fresh condition.
They would ideally serve the ends desired by Smith (1977:
123-125, 150-155) for "representative'' specimens. I have
discussed most of the processes of typification of sanction-
ed names elsewhere (Korf, 1982a, 1982b), and _ shall not
belabor the matter further here.
Since the type method requires suitable types, mycolo-
gists must now devote themselves (1) to finding holotypes
when they exist, (2) to designating lectotypes from undoubt-
bom or cial imacerial,. and, ‘far-.more frequently,” (3) to
providing adequate neotypes for old names that are now
"typified' by illustrations ('iconotypes') or by descrip-
tions alone.
First in importance will be specimen typification of
mames used in the two sanctioning works. Scrupulous care
must be used in designating neotypes, as the Code reminds
fis, (bulmesuch ‘care’ will result in neotypes (preferably
topotypes from the type locality) that will fix the names
as currently employed (at least in a great majority of
cases). Only later will we need to consider names placed in
synonymy by the sanctioning authors, or names overlooked or
ignored by them. When such an early name, after specimen
Eypiftication, ‘competes with a sanctioned: name, it will
merely fall into synonymy with the sanctioned (protected)
mane such a “name turns out: not. to’ *be)\ treated’ in’ the
sanctioning work, it also will need to be specimen typified
and will date from the date of its original publication.
mie principle of priority will’ in some cases then force us
to adopt an old, forgotten name instead of a name in use
but described later. This particular argument was advanced
acy a’ reason _for ‘not. abandoning later starting points, but
fPieprlority jis to be, served (Cand it ts °a-veuiding principle
of our Code) then it should be allowed. to operate. If
Quélet renamed. a species in the 1880's that Bulliard had
Previousiy described in’ 1/91,- why should we continue to
give credit to Quélet when Bulliard had come to the same
conclusion nearly a century earlier? Where such later spe-
fres senames are of J'major economic’ importance," we can
resort to species conservation. ?
SANCTIONED NAMES OR SANCTIONED EPITHETS?
It is unfortunate but true that legislators often make
Bemantic; errors. What zoologists ‘call)a *"species ‘name"’
metanists call.a.''species epithet.'' The various editions ‘of
Though nomina specifica conservanda were (at long last) permitted under the Code
adopted at Sydney in 1981, what botanists really intended to do, I remain convinced, was
to conserve species epithets, not species names. The Vice-Rapporteur (Greuter and Voss,
1982: 54-55.) explained that under the new provisions of Art. 14 what will be conserved
are basionyms. These are conserved against all homonyms and against all listed synonyms.
When the epithet of that conserved basionym is transferred, it retains its priority (at
the same rank) against any combination based on a rejected name. Horrendous circumlocu-
tions were introduced into the new wording of Art. 14 merely to avoid admitting that we
wanted to conserve species epithets rather than species names.
348
the ICBN have been plagued by use of 'name' in many places
where ‘'epithet' had been intended. One of the most flagrant
of these errors has been in regard to what) is. 'protected,
in Fries's Systema/Elenchus (and now in Persoon's Synopsis)
by the 1950-ff. Codes. Remember that the wording was (and
still is). "Names of FUNGI CAETERI ....’" What was incendee
was not only that names used by these authors were to be
preserved, but also epithets, as can best be shown in an
example:
Agaricus melleus Vahl was published in 1790, and is a
sanctioned name in Fries's Systema. One of the synonyms
cited by Fries is Agaricus obscurus Schaeff., published in
1762. With the new 1753 starting-point date, both names are
validly published, and were it not for the sanetioned
status accorded Vahl's name, we would be obliged to take up
Schaeffer's name on the basis of priority.* This is the
fungus known today as Armillaria mellea [or as Armillariella
mellea by those who (I agree with Watling et al., 1982) have
selected an ineligible type for the name Armillariajc slings
is indeed Vahl's name that was sanctioned by Fries, what
happens when we treat the epithet in another genus, Armilla-
ria or Armillariella? Neither Armillaria mellea nor Armilla-
riella mellea were names sanctioned by Fries,. and neither
would have priority status over the names "Armillaria obscu-
ra (Schaeff.) Author-X" or over "Armillariella obscura
(Schaeff£.) Author-Y." It was’ certainly never the intenetes
of those who. framed the wording in 1950 that gavey privae
ledged status to Fries's works nor those who adopted almost
identical wording in 1981 for providing sanctioned status
for Persoon's and Fries's works that old names synonymized
in the protected works could ever rise again to haunt ete
nomenclaturally! When they wrote ''Names of FUNGI CAETERI
..'' they clearly meant ''Names and epithets of FUNGI CAETERI
..'' were to be accorded sanctioned status. Not only @ame
the several generic and suprageneric names in the sanction-
ing works to have privileged status (at their rank), but
likewise epithets (infrageneric epithets, specific epithets,
and infraspecific epithets) have a priviledged status. When
such epithets are transferred (and retain their same rank)
it was clearly our intent that they should, retain’ theme
special status.
Another very important point needs to be made here:
sanctioned names and epithets are sanctioned against all
"synonyms and homonyms,'' whether these are mentioned in the
sanctioning work or not. In this regard sanctioned names
differ markedly from conserved names. Both take priority
over homonyms whether cited or not, but conserved names
take priority only over those earlier synonyms that are
specifically listed in the Appendices as nomina re jicienda>?
For our purposes here we shall assume that Fries was correct in synonymizing the two
names, and that they are indeed taxonomic synonyms.
Ever since first introduced, conserved family names have lacked listed nomina rejicien-
da despite the Codes' requirements that this be done! A new ‘housekeeping’ proposal was
introduced on the floor at Sydney (Greuter and Voss, 1982: 51-52) to exempt family names
349
inwour example» “Agaricus melleus Vahl. “:°Fr. as’ also pro-
tected against discovery of any pre-1790 name, even one not
eited@by Fries ,swhich weticonclude is a synonym. (And, as I
have “shown above, the intent is that the epithet 'mellea,
sus, "-Uum Vahl. : Fr." is similarly protected in Armillaria
and in any other genus to which it may be transferred at
species rank.)
RANKS OF SANCTIONED NAMES
Anyone who has worked with the sanctioning books will
be well aware that though genera and species are easily
recognized ranks, many of the suprageneric taxa bear ranks
unlike those in modern use, while many of the infrageneric
Categories are undesignated or are at times at ranks not
permited under the Code.® Infraspecific categories are
Similarly of several levels, often not indicated; following
tradition, "the rank of subspecies under species is marked
by lettere or figures; that of varieties by the series of
Sree lcapers, 8&5 (64 ¥. etc. Groups below Varieties “and
Nout preeds) (mule of (florists) are indicated) by letters,
Peres or typographical "signs,,) according to the williof
the author'' (de Candolle, 1868). (Art. 35 of the ICBN deals
in detail with cases where clear indication of rank is not
provided.) What mycologists must now keep in mind, of
Course, is that even if the rank of the taxon in one of the
sanctioning works is uncertain, the name is nonetheless
Sanctioned, and thus the sanctioning treatment should be
weighed when typifying a name (at whatever rank!). Unlike
priority, which is inoperative outside rank, typification
remains constant, even though rank may change. Should Agari-
cus melleus be reduced to subspecific or varietal rank
under another species name, it would retain the same type
Specimen it had when originally published at species rank
(or when neotypified at any rank).
SOME CARDINAL PRINCIPLES FOR
BS EPER Si) ANDO TSAR os CUTAT IONS
Having spent something over two years now working with
how to cite names under the new provisions adopted at
oydney, I have come to see several possible pitfalls.
Perhaps I can prevent others from repeating my mistakes.
The main difficulty has been in attempting to make these
notations do more than one job, in a mistaken sense that
economy of citation is an ideal. Once it is firmly entrench-
ed in one's mind that such notation only indicates sanction-
ing of the particular generic or suprageneric name or infra-
from the requirement to list nomina rejicienda. The Rapporteur-Général saw no problems
arising over the proposal. Yet a clear distinction was thereby made between the treatment
of conserved generic and specific names and those of families. The treatment now afforded
conserved family names comes closer to the priority aspects of sanctioned fungal names
than it does the priority aspects of conserved generic or specific names.
The pertinent rule of the Code (Art. 33.4, 33.5) makes an exception for Fries's taxa
termed "tribes" (tribus) in the Systema, "which are treated as validly published." No
other misplaced ranks are permitted.
O50
generic, specific, _or infraspecific epithet," then wWeewar
the notation system becomes unambiguous and easy to apply.
Principle 1
"a. Pers..""and., “: -Fr."" are notations, which may (pesues
out of author citations in non-taxonomic work, but should
normally be used in any taxonomic treatment, since they
convey information about priorability, typification, or both.
Principle 2
Sanctioning is independent of rank, and the notations
Mm; Pers.” and ."":"(Fr.'©: in no way indicate ‘the ranks. auemenee
a name or epithet was sanctioned. The purpose of the nota-
tion is to say: ''Beware: this name (epithet) was adopted in
a sanctioning work, and has special priority and typifica-
tion status."
Principle 3
Priority of sanctioned names and epithets extends only
within the rank assigned by the sanctioning author. (E.g.,
an epithet, sanctioned at varietal level, when raised to
species rank does not retain priority over an “earbier,
validly published but non-sanctioned species epithet. )
Principle 4
Typification is independent of rank, even when ranks
differ between original and sanctioning treatments. Typifica-
tion of sanctioned names and epithets must be viewed not
only on the basis of the protologue of the original author,
but also of all other elements referred to (directly Gon
indirectly) in the sanctioning work.
Principle 5
When’ the’ notation “s Pers.'""or “: Brivis used temo
always immediately follow the name of the original author.
Even though a_ sanctioning author may have sanctioned a
previous combination, it is critical that the notationgime
inseparable from the name of the original author, for the
two must be transferred together when the epithet itself
changes generic placement or rank. [E.g., Octospora applana-
ta Hedw. 1801 was transferred to Peziza in 1805 by Albertini
and Schweinitz. It might seem logical to cite the mames
sanctioned by Fries in 1822, as ''Peziza applanata (Hedw.)
A. & SS. : ¥Fr.,'' but this is illusory! When we transtem clus
species to yet another genus or give it another rank, it is
“applanata (Hedw. : Fr.) that we will want to) transter as
that workers will still know of its special priority and
typification status. The correct citation inv Pézizais ae
applanata (Hedw. : Fr.) A. & S.'"' If treated as a ‘species Gm
Octospora, it should be cited as "0.-applanata’ Hedw. . Fram
even eee Fries did not sanction it under that generi¢
name. /
Again I find here convincing evidence that it is epithets of species and infraspecific
551
Principle 6
When a new combination is made in one of the sanction-
ing works, the sanctioning author thereby simultaneously sanc-
tions the basionym and makes the transfer. His name should
appear twice. [E.g., citations such as "Daedalea_ striata
amen) eee. Ov. Der SUL ata (awartz. ¢. Fr.) Fra. 2. Fr.”
should be avoided for a first transfer of Boletus striatus
Swaruze(now B.. striatus Swartz :; Fr.). The correct format
MorecnisG is ‘Daedalea striata’ (Swartz :'Fr.) Fr." This, clear—
ly differentiates the two actions by Fries, transferral and
Panceroning. lo ask ‘ss. Fr.'"\tordo<two jobs .will lead to ¢om-
plications when the name is transferred again. Everything
within the parentheses will transfer, and nothing outside
the parentheses will do so.8]
Principle 7
When a new taxon is published in a sanctioning work,
it is necessary in taxonomic works to cite the sanctioning
author's name twice, separated by a colon. (E.g., ''Thelepho-
ra avellana Fr. .: Fr."' would be the preferred citation for
a new species published by Fries in 1821 in the Systema.
Not only, is -the name Fries's, but when ‘transferred it is
imperative thal its special priority status be recognized.
SOM Olgas 1t 16 “treated. at. the «rank of, ‘species, , the
epithet has priority over any epithet published earlier.)
Principle 8
When two competing names of the same date have both
been sanctioned, the first author to choose one over the
other should be followed, in accordance with the general
PeectiComorn IGEN (Art. 57.2). When they -are.of different
faves, spelerity of original publication will take effect
eM TNE (eat
taxa that are sanctioned. One could argue that in this instance both the name (combina-
tion) "P. applanata" and the epithet "applanata Hedw." were sanctioned, but it is only
critical that the sanctioning author's mame be connected to the epithet when it is
transferred, and of no importance that it be attached to the name (combination).
A citation such as Peziza bolaris Batsch : Fr. var. explanata (Holmsk. : Fr.) Fr.
conveys the information that (1) Holmskjold erected this taxon in some other genus or at
some other rank, or at the rank of varietas under some other species name, (2) that the
epithets bolaris and explanata are both sanctioned, and (3) that Fries (at some time)
transferred Holmskjold's epithet to varietal status under P. bolaris. Though the transfer
was actually made in the sanctioning work, the desire therefore to cite this as "P.
bolaris Batsch : Fr. var. explanata (Holmsk.) : Fr." must be resisted. While that would
convey the additional information that the transfer was made in the sanctioning work and
not in an earlier or later publication by Fries, such is not the purpose of ": Pers." and
": Fr." notations. Better we know what to transfer, "explanata (Holmsk. : Fr.),' when the
epithet is moved than particulars about when or where a combination was made, not a
normal function of author citations.
S52
ACKNOWLEDGEMENTS
I thank countless students and colleagues for stimulating discussion
and arguments that have brought me to my present understanding of the
Code, and to those several who have read and revised drafts of this
paper. Any faulty reasoning is my own responsibility, so I decline to
name them here lest their reputations be tarnished. Wherever I have
tripped, I welcome corrections and improvements. In the words of W.B.
Grove (1937: 367), "nomenclatorialism" is indeed fraught with "multitudi-
nous meticulosities."'
Pet BRATURE eit ED
CANDOLLE, A., de. 1868. Laws of Botanical Nomenclature Adopted
by the International Botanical Congress held at Paris in August, 1867;
together with an Historical Introduction and a Commentary. 72 pp. lL.
Reeve & Co., London.
COMPERE, P. OU Si Citation des auteurs antérieurs au point de
départ: ex ou crochets? Taxon 22: 702-703.
DEMOULIN, V., D.L. HAWKSWORTH, R.P. KORF & GZ. POUZAR. 1981. A
solution to the starting point problem in the nomenclature of fungi.
TaxoOnes0=) 52-05).
DONK, M.A. 1957. Typification and later starting-points. Taxon
G2, (245=256.
1961. The citation of authors of revalidated names.
Taxon 10: 66-69.
GREUTER, W. & E.G. VOSS. 1982. Report on botanical nomenclature
- Sydney 1981. Englera 2: 1-124.
GROVE, W.B. VO37 3 British Stem- and Leaf-fungi, vol. 2. 406
pp. Cambridge University Press, Cambridge.
INOIRUD AR GIP 1982a. Citation of authors' names and the typifica-
tion of names of fungal taxa published between 1753 and 1832 under the
changes in the Code of Nomenclature enacted in 1981. Mycologia 74:
259052 50%
1982b. Mycological and lichenological implications of
changes in the Code of Nomenclature enacted in 1981. Mycotaxon 14: -
476-490.
KORE Reb So eo. KOHN. 1979. Later starting point blues. I:
Monilia fructigena. Mycotaxon 9: 521-522.
& . 1980. Later starting point blues. II. Dumontinia
tuberosa: the thorny thickets of synonymy and some examples of nomencla-
forialrsm) “Mycotaxon 112) 381-395.
PETERSEN, R.H. 1975. Specific and infraspecific names for fungi
used in 1821. Part I. Introduction, A & B. Mycotaxon 1: 149-188.
SMIcnHG AH. EO Teles Speciation in Lactarius. pp. 1235255. In
Clémencon, H. [ed.], The Species Concept in Hymenomycetes. Bibliotheca
Mycologica 61. 444 pp. J. Cramer, Vaduz.
WATLING, R., G.A. KILE, & N.M. GREGORY. 1982. The genus Armilla-
ria - nomenclature, typification, the identity of Armillaria mellea and
species differentiation. Trans. Brit. Mycol. Soc. 78: 271-285.
MYCOTAXON
NiOde ANE NOU 2, pps, 355-579 January-March 1983
ON THE GENERA COCHLIOBOLUS AND PSEUDOCOCHLIOBOLUS
J.L. ALCORN
Plant Pathology Branch, Department of Primary Industries,
Indoorooptlly, Queensland, 4068, Australta
Abstract
Intraspecific variation in the two
characteristics used to distinguish Pseudo-
cochltobolus from Cochitobolus highlight the
difficulties involved in keeping these genera
separate, and it is proposed that the former
name be put into synonymy. The anamorphs of
Cochltobolus species are species of Bipolaris
and Curvularta, and these genera are compared.
Observations on conidium septation are reported,
and application of the terms '‘euseptate' and
"distoseptate' is discussed. The new combin-
ations Cochltobolus australtensts (Tsuda & Ueyama)
comb. nov., C. ntstkadot (Tsuda, Ueyama §&
Nishihara) comb. nov., and Btpolarts chloridts
(Alcorn) comb. nov. are proposed. In addition
the combination B. homomorphus (Luttrell §& Rogerson)
Subram. is validated, and a lectotype designated
for C. heterostrophus (Drechsler) Drechsler.
INTRODUCTION
When fungi in EZu-Helmtnthosporium (Btpolarts) were
first shown to have ascal state connections, those teleo-
morphs were described as species of Ophtobolus. In
describing 0. heterostrophus Drechsler, Drechsler (1925)
indicated that 'the leaf spot fungus is tentatively
assigned to Ophtobolus', and that this was 'not intended to
imply close relationship to certain species referred to this
genus' such as 0. cartcett (Berk. & Br.) Sacc. and
354
O. herpotrichus (Fr.) Sacc. Four more correlations between
Helmtnthosportum species and Ophtobolus were soon identified.
Ito and Kuribayashi (1927) described 0. mtyabeanus Ito §&
Kurib. as the perfect state of H. oryzae B. de Haan. The
other species described in this period were 0. kusanot
Nisik. (Nisikado 1928), 0. sativus Ito §& Kurib. (Ito §&
Kuribayashi in Kuribayashi 1929), and 0. setariae Ito §&
Kurib. (Ito §& Kuribayashi in Ito 1930). The anamorphs
recorded for these fungi were respectively H. kusanot Nisik.,
H. sativum Pamm., King & Bakke, and H. setartae Saw.
Cochliobolus was erected by Drechsler (1934) for the
fungus he had described previously as 0. heterostroplus.
The main features distinguishing the new genus from Ophtobolus
were greater ascus and ascospore width, the strongly helicoid
arrangement of spores in the ascus, and the associated
conidial state. Two species of Ophiobolus from Ambrosta
trifida L. were found to produce Phoma-like conidial states
in culture, while other species produced no anamorphs at all.
In contrast the anamorph associated with the type species of
Cochltobolus was a species of Helminthosportum exhibiting
bipolar germination (H. maydts Nisik. § Miyake). Drechsler
realized that several other species described in Ophtobolus
belonged in the new genus. He included 0. kusanot,
O. mtyabeanus, O. sativus, and 0. setartae in his concept
of Cochltobolus, but did not formally transfer the names
of these species. In addition to the four species just
listed, Drechsler regarded the unnamed Ophtobolus states of
H. micropus Drechsler and H. stenosptlwn Drechsler as
belonging in Coechltobolus. Transfer to Cochltoholus of the
names of the four Ophtobolus species accepted by Drechsler
was made by Dastur (1942) who also added C. trittcr Dastur.
Subsequently the genus has been expanded to include 32
species, many of which have Bipolarts anamorphs (Table HE ae
The correlation of some Btpolarts species with the
teleomorph genus Pseudocochltobolus (Tsuda, Ueyama, &
Nishihara 1977; Tsuda § Ueyama 1981) introduced hetero-
geneity into the anamorph-teleomorph connection previously
thought to be predictive, namely that all Btpolarts species
will prove to have Cochliobolus ascogenous states (Luttrell
1977, 1978). The differences between the two teleomorph
genera are examined below.
3555
Table 1. Cochltobolus species and their anamorphs
Species Author(s) Anamorph *
bicolor Paul & Parbery 1966 B. bicolor
boutelouae Sprague 1951 unknown
carbonum Nelson 1959 B. zeicola
chloridis Alcorn 1978 D. chloridis
cymbopogonis Hall §& Sivanesan 1972 C. cymbopogonis
cynodontis Nelson 1964a B. cynodontis
dactyloctenii Alcorn 1982 B. dactyloctenii
geniculatus Nelson 1964c C. geniculata
hawaiiensis Alcorn 1978 B. hawaiiensis
heterostrophus (Drechsler) Drechsler 1934 B. maydis
homomorphus Luttrell § Rogerson 1959 B. homomorphus
intermedius Nelson 1960b C. intermedia
kusanoi (Nisikado) Drechsler ex Dastur B. kusanoi
1942
lunatus Nelson §& Haasis 1964 C. lunata
melinidis Alcorn 1982 B. melinidis
miakei Hino & Katumoto 1966 unknown
miyabeanus (Ito §& Kuribayashi) Drechsler B. oryzae
ex Dastur 1942
nodulosus Luttrell 1957 B. nodulosa
palmivora Rao & Chaudhury 1964 unknown
peregianensis Alcorn 1982 B. peregianensis
perotidis Alcorn 1982 B. perotidis
ravenelii Alcorn 198la B. ravenelii
sasae Hino §& Katumoto 1960 unknown
sativus (Ito & Kuribayashi) Drechsler B. sorokiniana
ex Dastur 1942
setariae (Ito & Kuribayashi) Drechsler B. setariae
ex Dastur 1942
sitharamii Reddy 1976 unknown
spicifer Nelson 1964b B. spicifera
sporoboli Castellani 1951 Ho4sp;
stenospilus Matsumoto § Yamamoto 1936 B. stenospila
tripogonis Alcorn 198la B. tripogonis
critic. Dastur 1942 HS
victoriae Nelson 1960a B. victoriae
* B= Btpolarts, C = Curvularta, D = Dreehslera, H = Helmitnthosportun
(sensu lato)
356
OBSERVATIONS ON COCHLIOBOLUS AND PSEUDOCOCHLIOBOLUS
Tsuda et al. (1977) distinguished their genus
Pseudocochltobolus from Cochltobolus on two characteristics.
One was the presence of stromatic tissue below the ascocarp
body in Pseudocochltobolus. The other was the degree of
ascospore coiling in the ascus, pronounced in Cochltobolus
and slight or absent in Pseudocochitobolus.
Stromatic tissue
In the type species of Cochltobolus, C. heterostrophus
(Drechsler) Drechsler, there was no stroma developed in
association with ascocarps formed on leaves of Zea mays L.
incubated in a moist chamber (Drechsler 1925, 1934).
Numerous ascomata were produced during a study of Australian
races and mating types of this fungus, but in no case was a
conspicuous supporting stroma developed (Alcorn 1975). In
the current study, ascomata were produced by paired cultures
on Sachs' agar + maize leaf, and examined carefully for any
supporting stromatic tissue. None was seen. Stromata were
not induced by growing paired cultures on filter paper, maize
grain, wheat straw, or Chlorts gayana Kunth leaf (each on
Sachs' agar), although ascocarps formed on these substrates.
In contrast, ascomata of Pseudocochltobolus ntstkadot Tsuda,
Ueyama & Nishihara are borne on a stroma, consisting of a
short cylindrical base or a flattened crust in the substrate
(Tsudaet -al~ 1977+, pers?) obs..)),
In other species assigned to Cochltobolus, a sterile
stromatic base is sometimes present below the ascogenous
locule. These species include C. raveneltt Alcorn,
C. trtpogonts Alcorn (Alcorn 198la), C. perottdts Alcorn
(Alcorn 1982), C. Zunatus Nelson & Haasis (Nelson § Haasis
1964), C. geniculatus Nelson (Nelson 1964c), and C. sptetfer
Nelson (Nelson 1964b). I have shown that in C. cymbopogonts
J.A. Hall “G Sivan., this, characteristiciis strongly
influenced by the culture medium. A distinct stroma is
commonly produced on Sachs' agar + Z. mays leaf, but only
rarely formed on water agar + wheat straw (Alcorn 1981b).
The ascomata of Cochltobolus species are ascostromatic
ascocarps (Luttrell 1973), and in some species additional
basal stromatic tissue occurs. It seems undesirable to use
such a variable attribute as a generic criterion, and such
use in the context of the taxa under discussion is rejected.
Jo?
Ascospore coiling
Tsuda et al. (1977) claimed that the degree of asco-
spore coiling was pronounced in Cochltobolus, and slight or
absent in Pseudocochltobolus. This characteristic was
examined in fresh material of C. heterostrophus produced
by pairing compatible isolates on Sachs' agar + Z. mays leaf.
Ascocarps were crushed individually in lactofuchsin, and
each mature ascus categorized with respect to degree of
ascospore coiling. The following categories were used:-
(a) spores tightly coiled in a helical path over the
whole length, except sometimes very near the tip
where the spores tended to be parallel to the
long axis of the ascus.
(b) spores tightly coiled in the lower two thirds,
loosely coiled to straight in the upper third.
(c) spores loosely coiled for the whole length of
the ascus.
(d) spores looped in the lower half, coiled helically
in? the upper halt.
The number of asci in each category was counted in
Pesascocarpoatiaple 2)... (In) addition to these, there were
low numbers of asci in which the spores were tightly coiled
in the upper and lower parts, and loosely coiled to straight
in the median section; or looped irregularly for the whole
length. This characteristic is thus more variable than
suggested by Drechsler's account of the species (Drechsler
1°72 Jee Ouaby. che report, of Isuda, ct al. (1977),
Table 2. Ascospore coiling in asci of Cochltobolus
heterostrophus
Coiling category (see text)
(a) (b) (c) (d)
358
Ascospore arrangement in asci of P. ntstkadot, the
type species of Pseudocochltobolus, was described as
parallel to loosely coiled or very rarely coiled in a close
helix (Tsuda et al. 1977). Maximum lengths of asci and
ascospores were reported as 258 um and 288 um respectively.
In an isotype specimen (IMI 214643), 75 asci from a single
ascocarp were examined. Sixty asci had spores which were
straight to loosely coiled over much of the ascus length.
In most of these asci, spores were coiled more tightly for
20-30 um in the apical section. Some were coiled apically
and basally but were more or less straight in the median
section. In the other asci examined, ascospores were
classified as moderately to closely coiled over their whole
length. ‘These categories are broad, ‘but there isva continua
between spores straight and parallel, and spores coiled in
a close helix. An attempt was made to produce ascocarps of
P. ntstkadot under the same cultural conditions as
C. heterostrophus. Unfortunately two isolates supplied by
Dr. Tsuda (1021-3, 7-3-2) failed to form the teleomorph when
paired and a direct comparison was not possible.
Ascospores are loosely coiled in other species, for
example C. hawattensts Alcorn, C. peregtanensts Alcorn,
C. perottdts, C. trtpogonts, C. dactyloctenit Alcorn and
C. cymbopogonts. In the last-named species ascospores
commonly are straight and parallel for much of the ascus
length, although Hall and Sivanesan (1972) did not report
this fact. They described ascospores as 'closely spirally
coiled in the ascus', but gave maximum lengths of asci and
ascospores as 275 um and 300 um (rarely 420 um) respectively.
My observations on an isolate of C. cymbopogonts studied
by Hall and Sivanesan (IMI 130402) indicate that many asci
have spores only loosely coiled, or straight for much of
the ascus length. Species which have some ascospores
closely coiled for most of the ascus length also have
maximum ascospore length greater than maximum ascus length.
In contrast, where coiling is not pronounced the maximum
ascospore length may be equal to or less than that of asci.
Examples of the latter type are C. gentculatus (ascospore
maximum 270 um, ascus maximum 290 um), C. hawattensts
(190 um, 205 um), C. kusanot (Nisik.) Drechsler ex Dastur
(170 um, 170 um), C. Zunatus (270 um, 300 um), C. nodulosus
Luttrell (193 um, 193 um), C. peregianensts (200 um, 255 um),
CG. sptetfer* (240. um, 260 um),,.C.. sporobold BE. Castely.
Sao
(140 um, 140 um), and C. trtpogonts (275 um, 310 um). For
C. cymbopogonits, the corresponding values are 370 um and
340 um (pers. obs.).
Discussion
Intraspecific variation in the characteristics used
by Tsuda et al. (1977) to separate Pseudocochltobolus and
Cochltobolus throws doubt on their use as generic
differentials. I believe the evidence does not support
the retention of Pseudocochltobolus on the grounds offered,
and suggest that the name be put into synonymy with
Cochltobolus.
ANAMORPHS OF COCHLIOBOLUS
The description of Cochltobolus included details of
the anamorph, and Drechsler (1934) indicated that the new
genus was 'defined in both its sexual and asexual stages'.
By definition therefore, other Cochltobolus species should
have anamorphs congeneric with Btpolarts maydis (Nisik. §&
Miyake) Shoem. Drechsler recognized that species such as
Helminthosportum gentculatum Tracy & Earle differed from
others then referred to Helmtnthosporitum sensu lato, but
he did not exclude them from the genus. Subsequently
H. gentculatun and similar species have been segregated
in Curvularta (Boedijn 1933). This genus has been con-
sidered quite distinct from Bipolaris (Ellis 1966, 1971;
Groves & Skolko 1945; Putterill 1954), but the teleomorphs
discovered for members of both genera have been assigned
to Cochltobolus (Table 1). This would seem contrary to
Drechsler's circumscription of the genus, unless Curvularta
and Bipolarts are not distinct. Von Arx and Luttrell
(1979) have in fact suggested that Btpolarts should be made
a synonym of Curvularita. I have examined this proposal
and present results of my observations below.
The genera Curvularta and Bipolaris
The genus Curvularta was erected in 1933, with
C. lunata (Wakker) Boedijn as the type species (Boedijn
1933). Conidia of C. ltunata are obovoid, curved, 3-septate,
with the penultimate cell swollen, thicker walled, and
darker than the other cells. Btpolaris was established in
360
1959, with B. maydts as the type species (Shoemaker 1959).
Conidia of B. maydts are fusoid, curved, multiseptate,
with cells concolorous and lacking any disproportionate
swelling.
Ellis (1971) has provided standarized descriptions
for many dematiaceous hyphomycetes.
His descriptions
of Curvularta and Drechslera sensu lato (which includes
Btpolarts) reveal no differences in characteristics of the
colonies, mycelium, stromata, conidiogenous cells or
conidiophores. The genera are distinguished on the basis
of conidium morphology, summarized for each genus in the
following table. These data are drawn from the accounts
Dye ll Sal 966 sel O7ien 1976):
Conidium characteristics in the genera Curvularia and
Drechslera sensu lato
Curvularta
solitary or catenate
acropleurogenous
simple, often curved
clavate, ellipsoidal,
broadly fusiform, obovoid
or pyriform
pale or dark brown, often
with some cells paler
smooth or verrucose
hilum flush or protruding
septate
swollen median cell or cells
present
Drechslera
solitary or catenate
acropleurogenous
simple, straight or
curved
clavate, cylindrical,
ellipsoidal, fusiform
or obclavate
straw-coloured, pale to
dark brown or olivac-
eous brown, sometimes
with end, cells paler
mostly smooth, rarely
verruculose
hilum flush or protrud-
ing
pseudoseptate
swollen median cell or
cells absent
361
This comparison highlights the similarities between
these genera. Curved conidia occur in both, but in
Curvularta the curvature is often associated with a dispro-
portionately swollen median cell or cells, a characteristic
previously stressed by Ellis (1966). The conidia of many
species of Drechslera and Bipolarts have numerous septa,
although some have only a few. Examples of species with
the latter condition are B. aqustraltensts (M.B. Ellis)
Tsuda & Ueyama, D. papendorfit (van der Aa) M.B. Ellis,
B. spietfera (Bain.) Subram., D. biseptata (Sace. § Roum.)
Richardson §& Fraser, and B. ravenelizi (Curt.) Shoem. Most
Curvularta species have only three or four septa in mature
conidia.
mere 1S an inportant difference in the nature of
conidial septa, used by Ellis as a differentiating
character in his key to genera (Ellis 1971, p. 21, couplet
193). Conidia of Drechslera are distoseptate (= pseudo-
septate sensu Ellis), while those of Curvularta are septate.
Luttrell (1963) has pointed out that the term ‘pseudoseptate'
has been used in two senses by Ellis. In the context of
comparing Drechslera and Curvularta, it is clear that Ellis
has used the term in the sense of 'distoseptate’, and that
'septate' means 'euseptate' when applied to Curvularta
conidia.
Euseptate and distoseptate conidia
The terms euseptate and distoseptate were introduced
by Luttrell (1963). Euseptate conidia have a single wall
from which true septa form as inward extensions.
Sportdesmtum and Vakrabeejga (= Nakataea) were used as
examples of genera with such conidia by Luttrell. Disto-
septate conidia have at least two distinct walls. There
iseaytnin, ieid outer wall to which septa are not joined.
This outer wall encloses a thick, hyaline inner wall. The
distosepta delimiting cells of the conidium are formed by
the inner wall alone, and the outer wall does not parti-
cipate. In some fungi, for example Btpolarts sorokiniana
(Sacc.) Shoem., the outer wall of the conidium is brittle
and easily fractured by pressure. In such conidia the
internal structures often can be expelled intact, revealing
cells enclosed by the thick hyaline wall (Luttrell 1963).
362
Other species Luttrell used as examples of fungi with disto-
septate conidia were D. avenacea (Curt. ex Cooke) Shoem.,
Detghtontella torulosa (Syd.) M.B. Ellis, and Corynespora
cassittcola (Berk. & Curt.) Wei. In another paper, Luttrell
(1964) reported that conidia of B. maydis and Exserohtlum
turetcum (Pass.) Leonard §& Suggs were distoseptate, but
that this characteristic was difficult to demonstrate in
the former species because of the tough outer wall.
Observations on conidium septation in various genera
Conidia were mounted in 3% KOH and fractured by
tapping or pressing on the coverslip, then stained by
irrigation with 0.5% aqueous phloxine. This stained proto-
plasm a deep pink colour, and after some time hyaline walls
became pale pink. The term ‘wall' is used an the followmy
section in a broad sense, to describe ' a morphological
entity of apparent structural uniformity' (Sutton § Sandhu
1969 1%
Septation in Btpolaris, Drechslera and Exserohtlum
Conidia of B. maydts, the generic type, have two
distinct walls which can be demonstrated by the treatment
described above. Under pressure the thin pigmented outer
wall splits to release the hyaline contents... [he laces
sometimes emerge intact, and consist of a hyaline wall
enclosing the cells of the conidium (Fig. 2). As reporved
by Luttrell (1964), the outer wall in this species is
stronger than that in B. sorokintana, and more pressure
must be applied to achieve the same degree of fragmentation.
The inner wall often ruptured under these conditions. |
Only slight pressure is'necessary to break the outer
wall in conidia of B. sorokintana, and the contents often
can be expelled intact. A thick hyaline wall encloses
more or less globose cells with deeply-staining protoplasm
(Fig. 3). <A fine dark transverse line is often visible
between the cells. There is no evidence of torn attachment
points for septa when all structures internal to the brown
outer wall have been expelled.
363
Conidia of other Bipolarts species, and of Drechslera
and Exserohitlum species, were examined by the same method.
Species in which conidia were found to be distoseptate are
listed in Table 3. The brittle nature of the outer conidial
wall in B. soroktntana was not matched in all other species,
and some resembled B. maydts in having a wall much more
resistant to fracturing under pressure.
Luttrell (1963) raised the possibility that a third
wall existed in conidia of B. sorokintana, enclosing the
spherical protoplasts. Evidence was obtained to support
this suggestion. An inner hyaline wall was seen enclosing
protoplasts in some extruded conidial contents. This third
wall layer was also seen in conidia of B. tndica Rai,
Wadhwani & Tewari (Fig. 4), but in that species the inner
wall was thinner than in B. soroktntana. Electron micro-
scopy has led to differing opinions on wall structure in
conidia of the latter species. Cole (1973) reported that
the wall in developing conidia is two-layered, while Old
and Robertson (1969) and Mills (1970) recognized three
distinct layers. Subsequently a fourth layer was discovered
by Old and Robertson (1970).
Septation in Curvularta species
When conidia of C. lunata (the generic type) are
ruptured under pressure, the wall breaks at one or both
eidsmtOure case part Or all ot the’ contents. [he extruded
portion consists of a thick, hyaline wall ensheathing the
protoplasm which often is still divided into cells by
hyaline cross walls (Fig. 5). In other instances the proto-
plasm was continuous in the extruded portion. Often all
protoplasm was expelled, leaving the extruded wall material
as a more or less cylindrical tube equal to or greater in
length than the conidium (Fig. 7). Expelled protoplasts
sometimes retained their shape, but burst with the appli-
cation of greater pressure. The dark bands which accent-
uate the septa in conidia of this species usually remain
within the conidium, torn centrally but still attached to
the outer wall. The torn segments of the! septa are
directed towards the point of egress of the cell contents
(Fig. 5). Sometimes septa did not rupture, but bulged
towards the apex or base of the conidium without releasing
the cell contents. Occasionally one or more dark septa
were carried out of the conidium, and remained as
364
Table 3. Btpolarts, Drechslera and Exserohilum species in which conidia are distoseptate
Species
australiensis
australis
avenae
bicolor
biseptata
brizae
bromi
campanulata
catenaria
chloridis
clavata
coicis
crustacea
cylindrica
cynodontis
dactylidis
dactyloctenii
dematioidea
dictyoides
ellisii
erythrospila
fugax
gedarefensis
graminea
hawaliensis
holmii
homomorphus
incurvata
indica
iridis
kusanoi
leersil
maydis
melinidis
micropus
Isolate *
7750a
77126
14880
77106
PDDCC 6205
7855
DAOM 169239
WU.60/78
116-8
7670
77144c
GBS, 192.29
7-3-2
1021-3
1353
7835
7842a
DAOM 126766
7909
478
7838a
DAOM 155097
148
655
8021
DAOM 159452
7770
7795b
ATCC 13409
7783
20114
DAR 29777
CBS 7137 529
7704
22228
7837
8452
Species
monoceras
multiformis
neergaardii
nicotiae
nobleae
nodulosa
oryzae
ovariicola
panici-miliacei
papendorfii
pedicellatum
peregianensis
perotidis
phlei
poae
prolatum
ravenelii
rostratum
sacchari
setariae
siccans
sorghicola
sorokiniana
spicifera
stenospila
subpapendorfii
teres
tetrarrhenae
tripogonis
triseptata
tritici-repentis
tuberosa
turcicum
urochloae
victoriae
yamadae
zeicola
Isolate
7792a
1215
DAOM 154282
PC 2230
5063
20927
$401
7813
CBS 199.29
Ast
4951
7848
7905a
146
2879
Epa
7976
77162
20721
8562
5066
719%
7723
77144b
DAOM 147411
PGE22 en
1555
772Sa
77158
PDDCC 6120
21615
WU.51/78
77159
WIWAT
IMI 89977 (ii)
CBS» 202529
77107a
i
* Voucher specimens for all isolates used in this study are deposited in BRIP
365
distinctive dark bands across the expelled portion (Fig. 6).
These septa appeared to have been torn from the outer peri-
clinal wall intact. In face view these expelled accentuated
septa were circular, and had a small central pore.
Twenty-five other Curvularta species were examined,
and all were found to have conidium septum structure
Similar to that of C. lunata (Table 4). The variations in
morphology of the septa and extruded contents of conidia
noted for that species also occurred in some of the other
species examined (Figs. 8-11). Conidia of some species
were very difficult to crush, for example in C. verructformis
Agarwal & Sahni, C. uncinata Bugnicourt, and C. gentculata
(Tracy §& Earle) Boedijn. In these species each cell tends
Co release its contents separately from a rupture in the
Fateratewallealeayvang the septa intact, but a hyaline inner
wall can still be demonstrated. C. catenulata Reddy §&
Bilgrami and C. trtttct Kumar & Nema are included in Table
4 on the basis of observations on intact conidia. The inner
wall visible in some conidia, especially those with a
swollen median cell, could not be shown by crushing, to
which conidia of these two species are very resistant.
Septa in conidia of Curvularta species are therefore
double-walled. There is a thin central pigmented plate
which commonly remains attached to the periclinal wall after
the conidia are ruptured, and a thicker hyaline wall which
sometimes can be expelled intact. Published illustrations
of Curvularta conidia generally indicate only a single
outer wall to which septa are joined. Exceptions are
provided by some of the drawings of conidia of C. detghtonit
M.B. Ellis, C. borrertae (Viégas) M.B. Ellis, C. andropogonts
(Zimm.) Boedijn, C. tuberculata Jain, and C. robusta
Kilpatrick & Luttrell, where an inner wall to some or all
of the cells is shown (Ellis 1966, 1971). Similarly the
conidia of C. tschaemt McKenzie are shown as either single-
or double-walled (McKenzie 1981).
366
367
Septation in Btpolarts cotcts
Septa in conidia of B. cotcts (Nisik.) Shoem. are
commonly accentuated by a dark band in a manner similar to
that of Curvularta conidia. Three isolates of B. cdotcts
have been examined, namely CBS 192.29 authentic for the
name Helmtnthosportum cotcts Nisik., and isolates 7-3-2 and
1021-3 cited by Tsuda et a1. (1977). Septum structure in
conidia of these isolates is similar to that in conidia of
Curvularta species (Figs. 12, 13), and a proposal to accept
this fungus as a species of Curvularta could be advanced
using this evidence. The binomial C. cotcts E. Castell.
(Castellani 1955) would be available; I have examined the
type specimen (IMI 53422) and agree with Luttrell (1969)
that the name refers to the same fungus as described by
Nisikado. Characteristics other than those of septum
structure, however, point to Btpolarts as a more appro-
priate genus than Curvularta for this taxon. Thus,
curvature of conidia is not associated with a dispro-
portionate swelling of one or more cells, nor is there any
marked darkening of median cells.
Septation in Sportdesmtum and Nakataea
It was found that conidia of Curvularta species
ditter il septum structure, from species of Bipolarie. In
the former genus conidia are not euseptate as narrowly
defined by Luttrell (1963), who used Sportdesmtum tropicale
M.B. Ellis and Nakataea stgmotdea (Cav.) Hara as examples
of fungi where this condition applies. The form of
septation in these species has been examined for comparison
with that in Btpolarts and Curvularta.
Figs. 1-11. Septum structure in conidia of Bipolarts and
Curvularta spp. (1, water mount; 2-11, fractured in 3%
KOH vand¥stained with phloxine). 1:6 2. 8. maydts.
Srmeeoronitana. A. B.-tudiceq. Note thin inner wall
around protoplast of second cell from base. 5. C. lunata,
with ruptured dark septa retained internally. 6.C. lunata,
one dark septum expelled at apex; intact protoplast free
of inner hyaline wall at base. 7. C. lunata, inner wall
ruptured at apex and releasing protoplasm. 8. C. cymbo-
pogonts, lower septum bulged but not ruptured. 9. C. robusta,
lower two cells intact, upper three cells partly freed and
with dark septa intact. 10. C. rtchardtae, inner wall
intact. 11. C. rtchardiae, inner wall ruptured to release
protoplasts. Scale = ca. 10 um, all figures.
368
Table 4. Curvularta species in which conidia are
"euseptate'
Species Isolate Species Isolate
affinis MRL.1085 lunata 81151
andropogonis 19862b oryzae IMI 103899
brachyspora 8130 ovoidea MRL.1222
catenulata MRL.1175 pallescens MRL.1086
clavata OLZ5c prasadii MRL .1087
cymbopogonis IMI 130402 protuberata TABSC ALES
eragrostidis 81114b 16397
fallax IMI 102690 richardiae 7708
geniculata IMI 103902 robusta ATCC 16788
harveyi WA.227 senegalensis 8025
inaequalis MRL.1091 triftoliz 77169
intermedia 8013b CrLtiey MRL .1193
ischaemi PDDCC 6142 uncinata T/1534a
verruciformis © IMI 5252
ee
An authentic specimen of Sporidesmium troptcale
(IMI 41526, dupl. in BRIP) was examined.
species commonly rupture laterally.
Conidia-of tnis
The cell contents are
not enclosed in a second wall as occurs in Curvularia
conidia, and when released often assume a diffuse amorphous
shape.
The conidial septa rupture in a manner similar to
that described for Curvularta, however, with the torn
fragments directed towards the cell where pressure was
released by partial or complete expulsion of the proto-
plasm (Fig. 19).
Splitting laterally under pressure is also common
in conidia of Nakataea stgmotdea.
Again there is no sugg-
estion of a thick hyaline inner wall similar to that seen
in Curvularta conidia.
Under moderate pressure the larger
median cells usually rupture, with much more pressure
necessary to break the polar celts.
Septa break in the
same way as do those of Sportdesmium troptcale (Fig. 16).
Unlike the conidia of S. tropteale, those of N. stgmotdea
appear to be double-walled, at least in the median two cells.
There is often a small triangular space where the middle
septum meets the periclinal wall.
In some conidia this
Space is much larger, and it appears that the cells on
either side of the median septum have walls which are stru-
cturally distinct from the outer conidial wall (Fig. 17).
19
Figs. 12-20. Septum structure in conidia of Btpolarts,
Curvularta, Drechslera, Nakataea, and Sportdesmtum spp. (14,
water mount; others as for Figs. 2-11). 12. B. cotets,
ruptured dark septa internally, distal septum carried out
IO0
intact. 13. B. cotets, face view of expelled dark septum (arrow).
14.,C. protuberata, conidium from hyaline mutant (7788c).
Compare with Fig. 1. 15. C. protuberata, hyaline mutant. 16.
N. stgmotdea. 17. N. stgmotdea, detail of median septum,
showing triangular space where septum meets periclinal wall.
18. D. trtseptata, inner wall constricted at accentuated septum
19.85. -tropicale, 20. D. erythrospiia, ruptured dark. basal
septum (a), and distoseptate inner portion expelled from ape».
(b). The cap on this structure is the apical part of the dark
outer wall, freed during rupturing. Scale = ca. 6.5 um Fig. 17
10 um all other figures.
>
370
The septa delimiting the end cells appear to be formed by
a single wall, namely the inner wall surrounding each
median cell. The limitations imposed by light microscopy
prohibit an unequivocal statement about septation in this
species, but it may be similar to that described for
Setrtdtum (Roberts & Swart 1980), which has thinner-walled,
pater end*»celts:
Discussion
Cochltobolus as defined by the type species
C. heterostrophus has anamorphs in Btpolaris. The
discovery that some Curvularta species have teleomorphs
which apparently could be referred to Cochliobolus has
introduced a divergence from that principle. It has also
raised the question of whether Btpolarts and Curvularta
are distinct. Conidia of the former genus are distoseptate,
while those of Curvularta can be termed euseptate in a
broad sense, but not in the way used by Luttrell (1963)
when referring to Sportdesmtum troptcale. He defined
euseptate conidia as being ‘surrounded by a single wall
and have true septa formed as inward extensions of the
lateral walls'. In a glossary of new terms at the end of
the paper cited, a less restrictive definition than that
given above was included, that is - 'Euseptate: conidium
with true septa consisting of diaphragms merging peri-
pherally with the lateral walls'. The latter terminology
presumably would allow for the existence of a wall or walls
other than the lateral wall from which the septa origin-
ated. In this study, such a wall was demonstrated in
conidia of Curvularta species.
Mangenot and Reisinger (1976) recognized three
distinct categories of septal walls in conidia. In their
classification the septa are defined as outgrowths of a
Single wall (euseptation), or from a second inner wall
(distoseptation). A third type, called a double process
by Mangenot and Reisinger, involved both walls in the
formation of septa. Conidium septation in the Curvularta
species examined during the present study is referable to
this last type, rather than to euseptation as narrowly
defined, and perhaps could be referred to as 'secondarily
euseptate’.
Sit
Considerable intergeneric variation in conidium
septum structure can be demonstrated by simple manipulation
and observation by light microscopy. The results reported
above show that uniformity in septum structure occurs
within genera such as Btpolaris and Curvularta. They also
support the observations of Sutton (1969) that classif-
ication of septa into two categories might prove to be an
oversimplification. Roberts and Swart (1980) concluded
that the terms euseptate and distoseptate had not been
clarified by studies of conidium fine structure in
Pestalottopsts, Setrtdtum, and similar genera, and that
further ‘elucidation was necessary before they could be
used to indicate distinctions of taxonomic importance.
HOLOMORPH TAXONOMY FOR COCHLIOBOLUS
The genus Cochlitoholus has been used to accommodate
fungi with anamorphs in Btpolarts, and also fungi with
anamorphs in Curvularta. This apparent heterogeneity
depends on the recognition of these anamorph genera as
distinct. Von Arx and Luttrell (1979) have suggested that
Btpolarts should be made a synonym of Curvularta. If this
were done, all anamorphs of Coechltobolus would be in
Curvularta. I think it is preferable to maintain these
genera as distinct, especially in view of the evidence on
septum structure presented in this paper. If other conidium
characteristics are intermediate between those of Btpolaris
and Curvularta, then a consideration of septum structure
should allow taxa of otherwise uncertain affinities to be
placed in one or other of these genera.
If Curvularta is accepted, then the suitability
of Cochliobolus as a name for fungi with Curvularta
anamorphs must be considered. Drechsler (1934) defined the
genus Cochliobolus in terms of teleomorphic and anamorphic
forms, and used the concept that the conidial state was an
important criterion in helping distinguish it from
Ophtobolus. He showed that Ophtobolus species were con-
nected with Phoma-like anamorphs, and that C. heterostrophus
and congeneric species were connected with conidial forms
in Helminthosporium sensu lato (Btpolarts). By the same
reasoning, it might be argued that Cochltobolus species
with Curvularia anamorphs cannot be maintained in the same
genus as those with Btpolarts conidial states, and that a
separate ascomycete genus should be established for them.
Sr2
I once held such a view but now retreat from that position,
which emphasized differences in septum structure, to one in
which similarities in conidium morphology are given more
weight. Thus the wall and septum structure in conidia of
Btpolarts and Curvularta are regarded as fundamentally
alike. “There 2s a thin pigmented outer wall) and agthicker
hyaline inner wall which takes part in septum formation.
In Curvularta species the septa are accentuated by a dark
band which is attached to the outer periclinal wall. When
conidia are fractured under pressure, \this "secondary.
septum' commonly ruptures centrally but sometimes is
carried out of the conidium. If the latter occurs, the
similarity of the extruded portion to that produced when
Btpolarts conidia are treated in the same way is obvious.
Other features highlight the resemblances described
above. When Btpolarts conidia are mounted in water, the
inner wall is much less obvious than in lactophenol, and
the cells appear as overlapping circles (Fig. 1). They
have often been illustrated in this form by various authors.
Curvularta conidia mounted in water do not assume this
aspect, and the dark septa still appear to be united with
the lateral wall. An isolate of C. protuberata Nelson §&
Hodges used in these studies (7788c) gave rise to a hyaline
mutant in culture, which produced numerous conidia lacking
pigment. When such conidia are viewed in water mounts,
there is a striking similarity to distoseptate conidia of
Bitpolarts (Fig. 14). Crushing these hyaline conidia in
KOH also reveals their congruity with this coniditon
(Fig. 15). It may be noted that conidia of C. robusta were
described as distoseptate by Kilpatrick and Luttrell (1967).
Accentuated conidial septa are also found in species
which undoubtedly belong in Drechslera, for example
D. demattotdea (Bubak & Wrob. ) Subram. § Jain; D. biseptara;
D. trtseptata (Drechsler) Subram. & Jain, and
D. erythrosptla (Drechsler) Shoem. In conidia of the last-
named fungus, the accentuated basal septum often ruptures
as described for Curvularia, while a normal distoseptate
endospore is expelled from an apical split (Fig. 20).
Similar variation in a single conidium was noted in
C. tnaequalts (Shear) Boedijn. In D. trtseptata, the
internal wall of some conidia had become constricted at the
septa, clearly showing the darkened septum free from the
outer wall (Fig. 18).
573
Although Btpolaris and Curvularta are obviously
similar, I do not believe they should be united as has
been suggested (von Arx @ Luttrell 1979). Nor do I
believe, however, that the differences between them are
sufficient to warrant the erection of a separate genus to
accommodate the teleomorphs of Curvularta. This leaves as
the only option a maintenance of the status quo, accepting
a certain degree of heterogeneity within the anamorphs of
Cochltobholus species. This is not without precedent. For
example in Nectrita the anamorphs might be considered
variations on the theme of cylindrical phialide, in which
increasing complexity of arrangement of conidiogenous cells
can be recognized (Samuels § Rossman 1979).
New combinations
Earlier in this paper I proposed that the name
Pseudocochltobolus be put into synonymy with Cochltobolus.
The following changes then become necessary.
Cochltobolus nitstkadot (Tsuda, Ueyama § Nishihara)
comb. nov.
Pseudocochltobolus ntstkadot Tsuda, Ueyama §
Nishihara, Mycologia soo; lll7,. 1977.
Cochltoholus australtensts (Tsuda §& Ueyama) comb.
nov.
Pseudocochltioholus australtensts Tsuda §& Ueyama,
Mycologva®/57 292.1981.
I shall show in a subsequent paper that Btpolarts
and Drechslera should be maintained as separate taxa.
Drechslera chloridis Alcorn is the only Cochltobolus
anamorph to which a binomial has been applied (Table 1),
and which is now considered misplaced. The necessary new
combination is:
Btpolarts chlortdts (Alcorn) comb. nov.
Drechslera chlortdts Alcorn, Trans. Br. mycol.
S0C4 16/0 4148, (1976.
In addition, the combination B. homomorphus (Luttrell §
Rogerson) Subram. (Subramanian 1971) was not validly
published (Art. 33, basionym not indicated). This ommission
PeSucorrected, here:
374
Btpolarts homomorphus (Luttrell §& Rogerson) Subram.
ex Alcorn
Helmtnthosportum homomorphus Luttrell & Rogerson,
Mycologia <5)" 1955. 1959,
A LECTOTYPE FOR COCHLIOBOLUS HETEROSTROPAUS
A specimen labelled as the type was obtained from
BPI. The label details agreed with. those, cited by
Drechsler (1925):
Cochltobolus heterostrophus (Drechs.) Drechs. on
Zea mays
Santord, Flas\ssent. 22. 1925
Coll. Ac Foster
Det. Charles Drechsler
An inner packet contains approximately 24 leaves or
leaf segments. They are quite narrow, and very thin,
suggesting that they originated from seedlings. Numerous
lesions typical of infection by C. heterostrophus are
present on the leaves, but a thorough search of all the
material revealed no ascocarps, and only very scanty
conidial sporulation. Twenty-seven other collections from
BPI labelled C. heterostrophus were examined, but no
ascigerous state was detected. No slides from the type
showing the ascal state are preserved in BPI (M. Stransky,
pers. comm. 1977), and it therefore appeared that a neo-
type would have to be selected to typify C. heterostrophus.
Fortunately, ascal material identified by Drechsler is
available. In a paper in which the new genus
Pseudocochtltobolus was established, Tsuda et al. (1977)
referred to a specimen of C. heterostrophus from BPI in a
manner suggesting that ascal material had been seen. This
was confirmed in correspondence (Tsuda, pers. comm. 1978),
and subsequently the specimen was obtained from BPI. The
typed outer label bears the data:
Cochltobolus heterostrophus Drechsler on Zea mays
Washineron- D.C.) Oct... /,, 1924
Coll. & Det. Charles Drechsler
OPE
A handwritten inner label shows that the original name
given was Ophtobolus heterostrophus, that the host is
"Philippine corn leaf', and that it was incubated in a
damp chamber for 30 days beginning Oct. 7, 1924. This
is some of the material collected in the Philippines in
November 1921 and referred to by Drechsler in his account
of 0. heterostrophus (Drechsler 1925, p. 707). The
specimen consists of the contents of three moist chambers,
made up on 9 cm diam. filter papers. There are seven
leaf segments in all, each 6-7 cm long, darkened by fungal
colonies and bearing ascocarps visible without magnif-
ication. The ascocarps are black, scattered, amphigenous,
erumpent, mostly with a prominent cylindrical to conical
beak and often with brown setae or hyphae over the upper
half of the body but not on the neck. Three ascocarps
were examined. They were 460-520 um high x 360-400 um
diam., with the beak 140-160 um high, 90-110 um diam. at
the base and 60-100 um at the apex. Asci were obclavate
to cylindrical, with a short pedicel, and measured 132-
165 x 22.5-30 wm. Ascospores were not released from the
asci when the ascocarps were dissected in a drop of water,
sO measurements of length were not possible. They were
closely coiled in the ascus, hyaline to faint yellowish
brown, multiseptate, and 5-10 pm diam. As reported by
Tsuda et al. (1977), there was no evidence of stromatic
Crssue at tne base of the ascocarps. Conidia‘on the
specimen were similar to those on 28 other collections of
C. heterostrophus in BPI examined previously.
The specimen described above is designated as the
lectotype for Cochltobolus heterostrophus, the generic
type, in order that this important species may be properly
typified.
ACKNOWLEDGEMENTS
I thank the curators of BPI and IMI for their
courtesy in lending specimens. The following institutions
and individuals are acknowledged for assistance with gifts
or exchange of cultures, or for collecting fresh specimens:
American Type Culture Collection, Canadian National
Collection of Fungus Cultures, Centraalbureau voor
Schimmelcultures, A. Hagan, G.I. Johnson, W.J. Jooste,
Kew) Leonard, too... Luttrell), E.H.C. McKenzie, °J. Mouchacca,
Aw .sOnions, “LK. Price, R:G. Rees, J:E. Sheridan,
Moo isuda, §.J. Upsher, J, Walker and J. Webster.
376
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MYCOTAXON
Vols. XV Ig, INO sa25e pps 600-386 January-March 1983
TREMELLALES OF ISRAEL
N. BINYAMINI
Department of Botany,
The George S. Wise Faculty of Life Setences,
Tel Avtv Untiverstty,Tel Aviv, Israel.
ABSTRACT
Nine taxa of tremellaceous fungi growing
mainly with oak and pine trees, are described.
They were found in the Galilee and the central
parts of the country and eight of them are new
records for Israel.
INTRODUCTION
The Agaricales of Israel have already received attention
(Binyamini, 1977, 1980 and Moser, Binyamini & Avizohar-
Hershenzon, 1977), but information about the Tremellales of
Israel has, until now, been neglected. There is neither a
flora, nor are there monographic studies of the tremellaceous
fungi covering the study area around Israel. The nearest
area is Greece, as exemplified in the Fungus Host Index of
Pantidou (1973).
Owing to its geographical dry climate, Israel has a
relatively few tremellaceous resupinate fungi. This report
is concerned with 9 taxa, of which 8 are newly recorded for
Israel. Most of the collections upon which this report is
based were made during the period 1974-1981/2. Some of the
examined specimens show slight differences compared with
European material, possibly because of the dry climatic
conditions, and consequently, it was deemed worthwhile to
provide at least a short description of the species in our
collections.
381
One species listed has already been recorded by Rayss
& Reichert (1952). The systematics of the families and
genera follow Martin (1952) and Lowy (1971). The source
of each record is indicated at the end of the description
and the herbarium numbers are given in parentheses. The
colours specified in parentheses are taken from Kornerup
& Wanscher (1967).° Microscopic structures were observed
on fresh material. All collections cited have been
deposited in the herbarium of the Department of Botany,
Mycology Division, Tel Aviv University.
TREME LLACEAE
Extdta glandulosa Fr., Syst. Mycol. 2 : Pig sa
224.1882
Fruit body firm gelatinous, dark brownish, becoming
black, glistening when dry; hymenium smooth to tubercu-
late. Flesh blackish. Hyphae with clamp connections;
probasidia ovate or elliptical, 10-16 x 8-14 um, becoming
cruciate-septate, spores hyaline, allantoid, 10-15 x
4-5 um.
Habitat. On dry branch of Quercus, Mt. Carmel, Ha'arbaim
Wood, 9.2.81 (81p137), Upper Galilee, Bar'am Wood, 3.2.81
(81p460) , Hiram Wood, 3.2.81 (81p451, 81p452).
Our specimens conform well with the descriptions of Martin
(1952) “and = Lowy (1971).
Extdta nucleata (Schw.) Burt, Ann. Missouri Fig. °2
Botw Gard. co). fav 1. Lo2i.
Fruit body cerebriform, gelatinous, whitish becoming brown
with pinkish tinge to brown vinaceous with whitish
granules; hymenium smooth. Hyphae with clamp connections;
probasidia ovate, 10-12 x 6-8 um, cruciate-septate,
epibasidia subcylindrical up to 60 x 3.5-4.5 um, spores
hyaline, 8-11.5 x 4-4.5 um, cylindrical curved, germinating
by repetition.
Habitat. On dry branch of Quercus, Mt. Carmel, Bet Oren S.
29.12.80 (80p117, 80p151), Ha'arbaim Wood, 9.2.81 (81p138).
Upper Galilee, Bar'am Wood, 3.2.81 (81p130), on dry branch
of Calycotome villosa, Hiram Wood, 3.2.81 (81p139).
The white colour which changes through brown pinkish to brown
vinaceous is characteristic of our specimens, and conform
well with the descriptions of Martin (1952) and Lowy (1971).
S52
Extdtopsts effusa (Bref. ex Sacc.) A. Moller, Figsome
Bot. Mitth.. ‘Tropen 8 7 8z.31695
Fruit body soft gelatinous, greyish hyaline to nearly
chalk white; hymenium finely granulose, smooth < 100 um,
Tha CK Hyphae with clamp connections; dikarioparaphyses
subcylindrical branched, forming a thin layer above the
basidia; probasidia ovoid to subglobose, 10-15 x 8-10um,
clamps at the base, epibasidia four, up to 40 x 4 um,
spores cylindrical curved to allantoid, 12-15 x 4-5 um,
germinating by repetition.
Habitat. On dry branch of Quercus, Mt. Carmel,,Ha'
arbaim Wood, 9.2.81 (81p280, 81p281, 81p284, 81p389) .
On dry branch of Rhammus, Upper Galilee, Bar'am Wood,
B.2e0l (Slp4asiy.
The effused gelatinous greyish-white basidiome, with
branched dikaryoparaphyses and larged curved spores is
characteristic of our specimens. Spores are slightly
snaller than those described by Lowy (1971) and Wells &
Raitviir (1977), but close to those of Christiansen
(1959) and Reide (19 70}.. According to Wells (1961),
E. effusa is a synonym of £. grisea (Pers.) Bourd. &
Maire, but Wells & Raitviir (1977) agree that F. effusa
is very closely related to Z. grisea and that the dif-
ferences in substrate, colour of dry specimens and
Spore size differentiate the two taxa.
Sebactna tnerustans (Fr.) Tul., Jour. Linn. Fig. 4
Soc... Bot, fis is 330 sao ee
Fruit body fleshy coriaceous, up to 1 mm thick, cream-
yellow (443, 4C2, 4B2), margin slightly fibrillose.
Hyphae without clamp connections; probasidia subglob-
ose, becoming cruciate-septate, 15-20 x 11-13 um, epi-
basidia 30-75 x 2.5-3 um, spores ovate to oblong, with
lateral apiculus, 10.5-15 x 5.5-7 um, germinating by
repetition.
Habitat. On dry branch of Pinus. Ramot Menashe, Goara,
Zoeok Webwac lym
The resupinate, cream-yellow fruit body, and large, ovate
spores characterise our specimens; in these respects
the collection conforms well to the descriptions of
Martin (1952), Christiansen (1959) and Lowy (1971).
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Tremellavfimbriata Fr., Syst. Mycol. 2\.: 212, 1822.
=, jolttaced Pre; ovst., Mycol. 2 js12i2eeleec.
Fruit body leaf-like folds,gelatinous, cinnamon-brown,
becoming vinaceous black on drying. Hyphae with clamp
connections; probasidia subglobose, becoming cruciate-
septate, 12-18 x 10-15 um, epibasidia up to 70 um, spores
subglobose-ovate, 8-9 x 5-7 um, germinating by repetition.
Habitat. On dry branch of Quercus, Mt. Carmel, Bet Oren S.
20.1.81 (81p122).
Our specimens were slightly dry when found, but the micro-
scopic characters, such as clamped hyphae and spores,all
conform well to the descriptions of Martin (1952) and Lowy
(L971).
Tremella mesenterteca Fr., Syst. Mycol. 2 : 214 Fig. 5
1822.
=P oiutescens Fr., Syst. Mycol. *2 3121318227
Fruit body cerebriform to lobate, usually large, < 10 cm
in length, gelatinous, orange, or golden yellow. Hyphae
with clamp connections; probasidia subglobose, becoming
cruciate-septate, 10-17 x 8-14 um, epibasidia up to 75 um,
spores subglobose to ovoid, 10-14 x 8-12 um, germinating
by repetition.
Habitat. On dry branch of Quercus. Upper Galilee, Bar'am
Wood, 23.12.72. (72p152), (9.3.81) (81p209)) Mt.7 Carmel cet
Oren S., 29.12.80) (S0p118),0 87.78.82 (820103)F,
This, our commonest large Tremella, is frequently found on
dry branches and trunks of oak. Lowy (1971) supports the
interpretation of considering 7. mesenteritca Fr. and 7.
lutescens Fr. to be a single variable species.
DACRY MY CETACEAE
Daerymyces delitquescens (Merat) Duby, Bot. Gall.
PPT es PAS Te
Fruit body lenticular, up to 5 mm, gelatinous, orange-yellow
or pallid (4B6), becoming reddish brown on drying, smooth.
Hyphae with clamp connections, smooth or rarely roughened;
probasidia subclavate, 30-50 x 4-6 um, epibasidia bifurcate,
up to 15 x 3 um, spores allantoid, becoming 3 septate, 14-
16 x 5-6.5 um.
Boo
Habitat. On dry branch of Quercus, Mt. Carmel, Ha'arbaim Wood,
9.2.81 (81p135, 81p286). On dry branch of Pinus, Bet Oren,
9.2.81 (8ipl41,..81p169) .
The lenticular or sometimes flattened orange-yellow fruit
body, with bifurcate epibasidia and large allantoid septate
spores are characteristic for our specimens and conform well
to the description of Martin (1952).
-Calocera cornea (Batsch ex Fr.) Fr., strip. Fig. 6
Agri. Tems. 67. 1826.
Fruit body gelatinous to tough, narrowly club-shaped, un-
branched or very sparsely branched above, round in cross-
section, up to 15 mm high and 1 mm thick, pale yellow or
orange-yellow; hymenium smooth. Probasidia subclavate,
35-40 x 2.5-5 um, spores cylindrical, curved, 8-10 x 3-4.5 um.
Habitat. Scattered to gregarious, usually two to three
clustered together. Upper Galilee, Pa'ar Cave, on dry
branenwand trunk. of Quercus, 25.1.72 (72p505).,, 3.2281
(8ipl 22).
This small and beautiful species is variable in its colour,
size and degree of branching. The reddish tinge on drying,
which is described by Martin (1952) and Lowy (1971), is not
clear in our specimens.
AURICULARIACEAE
Auricularta auritcula(Hooker) Underwood in Barrett,
Mycologiay2a: 124 1910,
= Htrneola aurtcula-judae Berk., Outl. Brit. Fung.
2987 8860,
Habitat. On cut trunk of Ficus, Mique Izra‘'el 28.11.74
(74p106), 26.2.74 (74p104), 12.1.77 (77p113), Hulda, 8.2.75
(75p105), Ramat Hasharon, 1.9.78 (78p126).
This species has already been encountered in Israel and
described by Rayss and Reichert (1952). It has been found
several times; it is common each year.
386
LETERATURE CITED
Binyamini, N. 1977. Rare and interesting records of
Israeli agaric flora. Nova Hedwigia. 28
759-770.
Binyamini, N. 1980. Succession of Israel agaric flora.
Nova Hedwigia 32 : 185-198.
Christiansen, M. P. 1959. Danish Resupinate Fungi,
Part 1, Ascomycetes and Heterobasidiomycetes.
Dansk Botanisk Arkiv. 19(1) : 1-55.
Kornerup, A. & 3. H. Wanscher, 1967. Methuen Handbook of
Colour. Methuen & Co., Ltd.,London. 243 pp.
Lowy, B., 1971. Flora Neotropica Monograph No. 6.
Tremellales. Hafner Publishing Co., Inc., New
YOrK los) DD.
Martin, G. W. 1952. Revision of the North Central
Tremellales. State University of Iowa, Vol. 19
4233) 4s ele L22.
Moser, M., N. Binyamini & Z. Avizohar-Hershenzon. 1977.
New and noteworthy Russulales from Israel.
Trans. Br.uMycol. (SOG, 683). 2232 l=s/7 5
Pantidou, M. E.1973. Fungus Host Index for Greece.
Benaki Phytopathological Institute, Kiphissia,
Athens, 382 pp.
Rayss, T. and Reichert, I. 1952. Kemehim vepitriot
(Mushrooms) ,Tarshish, Jerusalem, 84 pp. (In
Hebrew) .
Reid, D. A. 1970. New or interesting records of British
Hymenomycetes, IV. Tran. Br. Mycol. Soc. 55(3):
413-441.
Wells, K. 1961. Studies af some Tremellaceae IV. Extdiopsts.
Mycologiay;yi53* 0317-370).
Wells, K. and A.Raitviir. 1977. The species of Exidtopsts
(Tremellaceae) of the U.S.S.R. Mycologia 69 (5)
987-1007.
MYCOTAXON
Vols XVIntNowe25 pia 58 /=595 January-March 1983
A NEW JAPANESE SPECIES OF NEOCOSMOSPORA FROM MARINE
SLUDGES
SETICHI UEDA! AND SHUN-ICHI UDAGAWA“
1 Nagasaki Prefectural Institute of Public Health and
Environmental Sciences, Nameshi l-chome, Nagasaki 852,
Japan
2 National Institute of Hygienic Sciences, Kamiyoga
l-chome, Setagaya-ku, Tokyo 158, Japan
mince: 1699 whenrl 4) F ..omith vestabiished the genus WNeo-
cosmospora with N. vasinfecta as the type, a total of 5
Species and 3 varieties has been described (Mahoney, 1976;
Donscnet ale, 1930) Only one of these; N. striata
Udagawa et Horie (1975), has ascospores covered with 8-10
transverse crests. A second member of Neocosmospora with
striate ascospores was isolated during a continuing survey
of the polluted fungi of Nagasaki Prefecture, Japan (Ueda,
Leo.) it this, paper a New species based’ on the isolate is
described and the major differences distinguishing the
species 7r0m N. striata are presented. [he specimen
studied is preserved at the Mycological Herbarium, National
Inseituce of Hygienic Serzences, Tokyo (NHE).
Neocosmospora tenuicristata Ueda et Udagawa, sp. nov.
(Figs. 1-12)
Stat. Anam. Acremonium tenuicristatum Ueda et Udagawa,
anam. sp. nov.
Coloniae in agaro cum decocto tuberorum et dextroso
celeriter crescentes, funiculosae, tenues, vinaceo-bubali-
nae; perithecia abunde producentia, persicina vel badia,
cum hyphis aeriis et conidiis albis obtecta; reversum brun-
neum vel brunneo-aurantiacum vel hinnuleum.
Perithecia superficialia, dispersa vel aggregata,
rubro-brunnea, ovoidea vel pyriformia, 335-415 x 265-320
um, glabra vel parce pilosa; pili hyalini vel dilute flavi,
non ramosi, septati, leves, breves, hyphoidei, 6-8 um diam;
perithecii collum breve, conicum plus minusve valde rubro-
brunneum, 65-80 x 80-115 um, glabrum; peridium membrana-
388
wid S
Neocosmospora tenuicristata, NHL 2911.
B. .perithecial anttiah. 8G. 5cUs a0.
E. Conidia-bearing structures and conidia.
Fiori
A. Perithecium.
Ascospores.
389
ceum, pseudoparenchymaticum, dilute flavo-aurantiacum,
saepe semitranslucens, 'textura angularis'; stratum exteri-
us e cellulis dilute salmoneis, incrassatis, angularibus,
10-32 x 10-22 um compositum; stratum interius e cellulis
hyalinis, tenuibus, complanatis, angularibus compositun;
periphyses hyalinae, filiformes, septatae, non ramosae, ad
basim 3-4 um diam, ad apicem rotundatae. Asci octospori,
hyalini, cylindracei, 90-110 x 13-15 um, superne rotundati
Velmaui1dualcumetruncatl, Sine poro.apicala instructi, ibre-
viter stipitati, in fasciculo basali dispositi. Ascosporae
uniseriatae, unicellulares, flavo-brunneae, late ellip-
SO1ceaemve! ecllipsoideac, 1575-17 x 10-115 um, suttulatae,
incrassatae, Sine poro germinali instructi, subtiliter et
transverse striatae.
Mycelium e hyphis hyalinis vel dilute flavo-brunneis,
ramosis, septatis, levibus, 1.5-10 wm diam, saepe in funi-
culo aggregatis compositum. Conidiophora brevia, plerumque
15-35 x 3-5 um, hyalina, non ramosa vel pauci-ramosa,
HeViagmeseptataserecta,. ¢ mycelio vegetativo vel funiculis
hypharum et hyphis aeriis orientia, mononemata. Cellulae
conidiogenae monophialidicae vel polyphialidicae, plerumque
simplices sed interdum sympodice proliferae vel percur-
rentes, terminales, saepe in conidiophoris incorporatae,
rectae, hyalinae, 30-50(-125) x 2.5-4 um, superne gradatim
attenuatae, collari parvo praedita. Conidia hyalina, gut-
tulata, 0-l-septata, ellipsoideo-cylindracea vel fusi-
formia, parum incurvata, 6.5-15(-19) x 2.5-4.5 um, levia,
utrinque rotundata, in massa mucida aggregata. Chlamydo-
sporae absunt.
Holotypus: cultura NHL 2911 ex limo (marine sludge),
Nagasaki in Japonia, in’ 26,1.1981,; aS. Ueda, isolata. In
collectione fungorum 'National Institute of Hygienic Sci-
encess (NHL) .Tokyo, Japan".
Etymology: lat. tenui- = thin and cristatus = crested,
referring to the nature of the ascospore ornamentation.
Colonies on potato-dextrose agar spreading broadly,
attaining a diameter of 8.0 cm in 2 weeks at 23 C, funicu-
lose, consisting of a thin mycelial felt, Vinaceous Buff
(Rayner, 1970) in color; perithecia abundantly produced on
the felt, Peach or Bay (Rayner, 1970), covered loosely with
white aerial hyphae and conidia; reverse brown to brownish
orange or Fawn (Rayner, 1970).
Perithecia superficial, scattered or aggregated, at
first flesh-colored, then becoming pale pink or orange-red,
finally reddish brown, ovoid to pyriform, 335-415 x 265-320
um, glabrous or sparingly covered with hyaline to pale yel-
low, unbranched, septate, smooth-walled, short, hyphal-like
390
SOL
hairs measuring 6-8 wm in diam; neck short, conical, more
or less dark reddish brown, 65-80 x 80-115 um, glabrous, at
Ules0st10re region’ composed of short, cylindric cells;
peridium membraneous, pseudoparenchymatous, pale yellowish
orange, often semitransparent, textura angularis; outer
layer composed of pale salmon-colored, thick-walled, angu-
lar cells measuring 10-32 x 10-22 um; inner layer of hya-
line, thin-walled, flatten, angular cells; periphyses hya-
line, filiform, septate, unbranched, 3-4 um in diam at the
base, tapering to a rounded tip. Asci 8-spored, hyaline,
cylindric, 90-110 x13-15 um, rounded or somewhat truncate
above, without distinct apical structure, short-stalked
(stipe: up to 20 um long), borne in a basal fascicle. Asco-
spores uniseriate, one-celled, yellowish brown, broadly
eliipeem to ellipsoid, 13.5-17 x 10-11.5 um, guttulate,
thick-walled, without germ pore, with surface ornamented by
very fine ridges which are roughly transverse and wavy in
appearance, extruded at maturity in the form of a gelatin-
OUSmides ~epespore: ca. 1.5, um ‘thick.
Mycelium composed of hyaline to pale yellowish brown,
branched, septate, smooth-walled hyphae 1.5-10 um wide,
OMleneaveresatecd in funicles. .Perithecial initials arising
as) coiled or contorted side branches of hypha, then becom-
ing surrounded by hyphae arising from the neighboring
cells. Conidiophores short, usually 15-35 x 3-5 um, hya-
line, unbranched or a few branched, smooth-walled, septate,
arising directly from the vegetative mycelium or hyphal
ropes and aerial hyphae, mononematous. Conidiogenous
cells monophialidic or polyphialidic, usually simple but
sometimes proliferating sympodially or percurrent, termi-
nal, integrated, straight, hyaline, 30-50(-125) x 2.5-4 un,
tapering gradually to the minute apical collarette measur-
ing 2-2,5 * 1.5-2 pm; phialides sometimes reduced to a col-
fanette,onethehypha. .Conidia hyaline, guttulate, 0-1-
septate, ellipsoid-cylindric or fusiform, slightly incurv-
ed, 6.5-15(-19) x 2.5-4.5 um, smooth-walled, rounded at
both ends, aggregated in a slimy mass measuring 7.5-18 um
in diam. Chlamydospores absent.
At] 37 Coycrowth is slightly slower that at 23 C and
perithecial production is reduced.
Specimen examined: an isolate from marine sludge, at
Oomura Bay, Higashisonogi-gun, Nagasaki-pref., Japan, Jan.
Bios siz-o. 9. tenuicristata, NHL 2911.
2. Asci. 3. Ascospores, showing germination. 4. Perithe-
cial initial. 5-8. Conidia-bearing structures and conidia.
The line in all figures equals 20 um.
392
593
Zot Jol. >. Ueda, NIL 2911, holotype. Subculture of NHL
2911 has been deposited with the American Type Culture Col-
lection (ATCC).
Neocosmospora tenuicristata, with its striate asco-
spores and polyphialidic anamorph, is unique in its genus.
In the ascospore ornamentation, it has a strong resemblance
to N. striata Udagawa et Horie (1975), which differs by
having smaller perithecia with a cylindric neck, irregular-
elongate cells of the peridium in place of angular-shaped
cells, smaller asci and ascospores which are ornamented
with more rugged 8-10 transverse crests, and the absence of
anamorph. According to van Warmelo's SEM study (1976), the
patterns of surface ridges of ascospores in N. vasinfecta
and N. africana von Arx appeared to be quite similar. The
ascospore ornamentation in both species consists of a net-
work of anastomosing ridges over the surface, while the
ridges in N. tenuicristata are clearly separate as revealed
by SEM observation (Figs. 10-12) as well as examination
with the light microscope.
Another Interesting point in N. tenuicristata 1S its
anamorphic characters. there are a few species such as
A. hyalinulum (Sacc.) W. Gams and A. brunnescens W. Gams in
the form-genus Acremonium (Gams, 1971, 1975) having proli-
feration of phialides (polyphialides). Both Acremonium are
distinguished from the anamorph of N. tenuicristata by
their small (not exceeding 7 um in length and 2.5 um in
width) and catenated conidia, as well as by very slow-
growing fine mycelia. For the anamorph of N. vasinfecta,
Domsch et al. (1980) stated that ''Cephalosporium-like, but
as the colonies grow fast and the hyphae are wide, it
recalls the micro-conidial states of Fusarium rather than
Acremonium."' In fact, the growth-rate on usual media, and
the features of vegetative mycelium and conidiogenous
structures in N. tenuicristata are similar to those of Fu-
sarium proliferatum (Matsushima) Nirenberg in the sect.
Liseola. Matsushima (1971) firstly described Cephalospori-
um proliferatum as follows: Phialophora ex hyphis repenti-
bus vel aeriis ascendentia, simplicia vel ramosa, usque ad
100 um alta, 2-4 um crassa, rami terminales fertilia, sym-
Figs. 9-ll. N. tenuicristata, NHL 2911, and Fig. 12.
N. africana von Arx, No. 78-S-35-10.
9, Ascospores with light microscopy. 10-12. Ascospores
with scanning electron microscopy, showing their difference
in surface ornamentation. The line in Fig. 9 represents 20
um; in Figs. 10-12 equals 10 um.
394
podialiter sub apice proliferascentia et polyphialidibus
geniculatis formata. Conidia of C. proliferatum are more
or less clavate, one-celled, 5-11 x 2-4 um, and with a
truncate base. Matsushima failed to observe the macroconi-
dia on his material of C. proliferatum, but after comparing
with 46 isolates of Fusarium moniliforme Sheld. sensu
Wollenw. et Reinking pr. p., Nirenberg (1976) conctuded
that pyriform conidia of both are conspecific and transfer-
red C. proliferatum to the genus Fusarium. The anamorph of
N. tenuicristata differs from F. proliferatum in having
broader mycelium, larger ellipsoid-fusiform conidia with
rounded ends and in the absence of macroconidia.
Since the conidia of the previous Neocosmospora
species have usually been placed in Acremonium or Acremo-
nium-like genera, the anamorph of N. tenuicristata seems to
be better treated as Acremonium until the discovery of its
associated macroconidia. However, the occurrence of poly-
phialidic structures further emphasizes the affinity of
anamorphs of this and other Neocosmospora to Fusarium.
ACKNOWLEDGMENT
Appreciation is expressed to Dr. S-C. Jong, American
Type Culture Collection, for reviewing the manuscript.
LITERATURE CITED
Domsch, K. H., W. Gams, and T.-H. Anderson. 1980. Compen-
diumsof Soil Fungi. Vol. -1., pp. 509-sl0) = Academic
Press. London:
Gams, W. 1971. Cephalosporium-artige Schimmelpilze (Hy-
phomycetes)” Gustav Fischer, Stuttgart. 2625p.
Gams, W. 1975. Cephalosporium-like Hyphomycetes: Some
tropical ‘species. Trans. Br. mycolz js0G¢. (64. sc7=40 4
Mahoney, D. P. 1976. A new Neocosmospora from Galapagos
Island soil. Mycologia 68: 1111-1116.
Matsushima, T. 1971. Microfungi of the Solomon Islands
and Papua-New Guinea. Kobe. p. 11 and fig. 121-2.
Nirenberg, H. I. 1976. Untersuchungen Uber die morpholo-
gische und biologische Differenzierung in der Fusari-
um-Sektion Liseola. Mitt. Biol. Bundesanst. Land-
Forstwirtsch. Berlin-Dahlem 169: 1-117.
Rayner, Ry W/ 1970. A mycological colour chart. 9 Cramer.
§& British Mycological Society.
Udagawa, S., and Y. Horie. 1975. “Notes on some Japanese
Ascomycetes XIV. Trans. mycol. Soc. Japan 16: 337-
347.
395
Ueda, S. 1981. A mycofloral study of marine sludges in
Nagasaki, Japan. Proc. 25th Ann. Meeting Mycol. Soc.
Japan. (Pp. fo.
Warmelo, k. 1. van. 19760, Scanning electron microscopy of
Neocosmospora ascospores. Mycologia 68: 1181-1187.
MYCOTAXON
Vol “AVA, “No. 2, spp 396-402 January-March 1983
NEW RECORDS OF HYPOGEOUS ASCOMYCETES
IN ARIZONA
JACK S. STATES
Department of Biological Sciences
Northern Arizona University, Flagstaff, AZ 86011, USA
SUMMARY
The following hypogeous ascomycetes are
reported as new records for Arizona, USA:
Balsamia magnata, Elaphomyces verrucosus, E.
granulatus, Geopora cooperi f. cooperi, G.
cooperi f. gilkeyae, Tuber dryophilum, and T.
levissimum. All species are putative mycorrhizal
symbionts with Pinus ponderosa Laws. They were
major food items in the diet of the tassel-eared
squirrel, Sciurus aberti.
A nearly pure stand of Pinus ponderosa Laws. covers
7.5 million acres in northern Arizona and western New
Mexico. The tassel-eared squirrel, Sciurus' aberti
Woodhouse, and its various subspecies (Hofmeister and
Diersing, 1978), are closely associated with this
conifer. Citing a close dependence on ponderosa pine for
food and shelter, Keith (1965) and Hall (1981) point out a
restricted distribution of tassel-eared squirrels to the
southwestern United States and northern Mexico, with
particular abundance in Arizona and New Mexico.
Stephenson (1975) in a study of the food habits of
tassel-eared squirrels in northern Arizona, noted that
fungi are a major component in their diet throughout most
of the year, but he did not attempt to identify the
species involved. Upon examination of the spores in the
stomach contents provided by Stephenson, I determined that
the majority of the fungi consumed were hypogeous species,
both ascomycetes (truffles) and basidiomycetes (false
truffles). The ascomycete spores were particularly
597
interesting since they represented the genera Geopora,
Elaphomyces, and Tuber, all previously unreported from
Arizona. Although the squirrels proved to be much more
efficient truffle collectors, I have discovered, through
my own forays over the past three years, the fruitbodies
of the major ascomycete species whose spores are most
frequently represented in the squirrel diet.
The hypogeous fungi are little known in the
southwestern United States and only occasional collections
have been reported, especially in Smith et al. (1981).
Fogel and Trappe (1976) have reported the occurrence of
hypogeous ascomycetes in the Rocky Mountain region. Some
of these are quite similar to those reported here.
Nonetheless, I am able to add three additional species in
two genera as well as additional host tree associations as
presumptive ectomycorrhizae. All species described are
new records for Arizona. Collections are deposited in the
Deaver Herbarium, Northern Arizona University, and in the
herbarium of Oregon State University (OSC).
1. BALSAMIA MAGNATA Harkn. Sensu Trappe, Mycotaxon 2:119.
197 5%
A single collection of three ascocarps match in most
respects the description by Gilkey (1916) as Pseudobal-
samia magnata (Hark.) Gilkey. The very distinctive
brownish pink to reddish brown ascocarps measuring 0.5-1.5
cm in diameter made them difficult to detect among the
reddish jasper and chert debris of the mineral soil. The
verrucae and the occasional small surface depressions of
the peridium strongly resembled the surface texture of a
russett potato.
ASCOCARPS hypogeous in shallow mineral soil or only
slightly imbedded beneath a thin needle layer of Pinus
ponderosa. Basal tuft and surface opening canals (venae
externae) were not clearly evident.
COLLECTION EXAMINED: Coconino County--Jacob Lake,
June, elev. 2250 m., States AHF/1.
Ine, ELAPHOMYCES VERRUCOSUS Dodge, Ann. Mycol. 27:171-
LIZ 28 1929).
Species of Elaphomyces are difficult to identify.
Spore ornamentation as a prime character has been shown to
398
be variable according to conditions of desiccation and
developmental stage (Hawker, 1968). The separation of E.
verrucosus from related species is based primarily on
coloration of a marbled peridium when cut in cross
section. I have found peridial marbling and _ the
development of surface warts to be inconsistent when
comparing different collections. Trappe considers E&.
verrucosus as a synonym of E. decipiens (pers. comm.).
Until better resolution of this problem is achieved I am
satisfied with the present disposition of the
collections. The description also fits my collections
made in ponderosa pine forests in the Wyoming Black Hills,
States WHF14 and WHF32.
ASCOCARPS sphaerical to ovoid, variously depressed
when formed near rocky debris; the surface covered by a
dense, earthy crust easily separable and composed of Pinus
ponderosa mycorrhizae and brownish yellow mycelium;
peridium yellow or _ ochraceous, drying yellow brown,
verrucose with pointed to rounded yellow warts 90-200 Um
high and 100-175 Um wide, hyphal layers strongly to
inconspicuously marbled in cross section, when dry 2.0-3.0
Um thick, vinaceous brown with dull rose or rose gray
veins interspersed, the layer above the gleba light rose
tan; gleba powdery, dark brown to fuliginous; spores
globose, 22-28 (31) Um in diameter, ornamentation evenly
distributed and densely echinulate, friable after
desiccation and easily splitting away from the _ spore
surface when pressed; odor slight or not detectable.
COLLECTIONS EXAMINED: Coconino County--Jacob Lake,
June through August, elev. 2250 m., States AHF88, AHF89,
AHF117, AHF146, AHF209; Paradise Rd. Flagstaff, States
AHF149; Woody Mountain, States AHF103.
3. ELAPHOMYCES GRANULATUS Fr., Syst. Myc. 3:58. 1829.
Fogel and Trappe (1976) described FE. granulatus
collected in Colorado among ectomycorrhizae of Pinus
contorta. The characteristics of our specimen generally
follow theirs but some of the special features are listed
below.
ASCOCARPS beneath Pinus ponderosa litter, deep in
mineral soil, associated with a removable soil crust
containing pine ectomycorrhizae; peridium smooth, pallid
to tawny yellow with faint rusty brown on drying with
warts closely appressed as superficial black spots, 3-4 mm
pe)
thick when fresh and 2-3 mm when dry, tough but
cartilaginous and flexible, somewhat viscid when wet, the
outer layer in cross section grayish yellow with alternate
zones or lenses of brown hyphae in different shades below,
the light gray to tan layers above the gleba turn black
when cut or bruised; gleba gray brown to dark brown or
black at maturity, separating from peridial hyphae as a
central powdery mass of spores and hyphae; spores held in
globose clusters of eight by an evanescent ascus, globose,
12-22 wm in diameter including ornamentation, the smaller
spores dark brown, the larger spores brownish gray to
gray, ornamented with crowded spines joined to form small
groups of parallel ridges traversing the spores in an
undulating pattern, the fragile epispore splitting to
reveal a large sphaerical oil droplet imbedded in a
gelatinous matrix.
COLLECTIONS EXAMINED: Coconino County--Jacob Lake,
June, elev. 2250 m., Lanphear AHF82; Woody Mountain,
August, elev. 2190 m., STATES AHF250.
A. GEOPORA COOPERT Harkn. f. COOPERI, Bull. Calif. Acad.
Seige lt 166.:1685:.
This species is likely to be widely distributed in
Arizona coniferous forests as indicated by its frequent
occurrence in fecal wastes of tassel-eared squirrels.
Fogel and Trappe (1976) recorded it from Colorado and I
have found it in the Medicine Bow Mountains of Wyoming,
States WHF31.
ASCOCARPS hypogeous among ectomycorrhizae of Pinus
ponderosa, Picea angelmanii, and mixed stands of pine and
Quercus gambelli. Mostly in exposed sites in rocky
mineral soil, May to November, elev. 2100-3300 m.
COLLECTIONS EXAMINED: Coconino County--White Horse
Hills, States AHF4; San Francisco Peaks, Snow Bowl ski
area, States AHF131; Casner’ Park, States AHFI174;
Transition Zone Horticultural Institute near Flagstaff,
States AHF175; Hoffderker Hills, States AHF105.
5. GEOPORA COOPERI Harkn. f.GILKEYAE Burdsall, Mycologia
ASCOCARPS were found to be gregarious in deep mineral
soil beneath Pinus ponderosa and Pseudotsuga menziesii
400
litter and surrounded by ectomycorrhizae. The collections
fit Burdsall’s description who reported it from Colorado
and Idaho. I have made collections of both forms of
Geopora in adjacent locations. Spore size is a constant
difference and only in f. GILKEYAE was the sporocarp odor
pronounced, much like fermented cider. Im addition, tiv
GILKEYAE possessed a cream to yellowish tan hymenium
whereas the hymenium of f. COOPERI was more typically
white to light cream in mature specimens.
COLLECTIONS EXAMINED: Coconino County—-—-Woody
Mountain, October to November, elev. 2280 m., Lanphear
AHF175; Paradise Rd., Flagstaff, States AHF178; Pinal
County--Santa Catalina Mountains, September, elev. 2440
m., States AHF242.
6. TUBER DRYOPHILUM Tul. Fungi Hypog. 147. 1851.
The discovery of Tuber in Arizona is noteworthy
because it represents a significant range extension for T.
dryophilum as well as T. levissimum recorded below. A
comparison of their striking spore ornamentation is
illustrated in Figure l.
ASCOCARPS hypogeous in mineral soil, 2-6 cm deep with
a moderate covering of leaf litter, primordia were noted
in mid-August and mature fruitbodies were noted in early
December, widely scattered among ectomycorrhizae of Pinus
ponderosa and Quercus gambellii, most abundant on steep
south-facing slopes in dense forest stands, rare
occurrences were noted beneath alligator juniper
(Juniperus depeanna), small, 0.5-1.5 cm in diameter, white
becoming yellow brown, irregularly subglobose, lightly
lobed with white furrows beneath the lobes; peridium
variable in thickness, 150-300 (400) wm thick; asci short
stipitate when immature, subglobose and persistent as the
spores develop, 1-4 rarely 5 spores per ascus; spores
globose-ellipsoid to nearly globose, 20-40 x 22-40 (45)
um , bearing prominent alveoli, the ornamentations
extending well beyond the spore surface, 3-5 (6) alveoli
spanning the greatest diameter, walls and ornamentations
becoming rich brown in age.
COLLECTIONS EXAMINED: Yavapai County--Five-mile
lake, August, elev. 2150 m., States AHF121; Coconino
County--Oak Creek Canyon, August to October, elev. 2080 m.
States AHF148, AHF167; Paradise Rd., Flagstaff, August to
401
December, elev. 2150 m, States AHF162, AHF180.
EG TUBER LEVISSIMUM Gilkey, Univ. Calif. Publ. Bot
Geo lo, 1916.
ASCOCARPS. O'.5-1.5° (220)' "em" in diameter, glabrous,
smooth texture, bruising tan to light brown, white
becoming light yellow brown at maturity, lined with white
furrows which extend to the interior as venae externae,
contours lobed especially in larger specimens; peridium
thine Drown plemented, 100-250 ‘tim thick, distinctly
pseudoparenchymatous becoming hyphal near _ the glebal
cavities; gleba in cross section marbled with tan to gray
brown fertile regions contrasting with the white venae;
asci subglobose becoming evanescent at maturity with 1-4
dark brown spores; spores subglobose to broadly ellipsoid,
alveolate with 5-6 (7) alveoli spanning the diameter of
the largest spores, small spores mostly subglobose, 44-53
(62) x 35-45 um, 3-4 alveoli across the diameter, taste
mild and richly mushroom-like.
COLLECTIONS EXAMINED: Coconino County--Paradise Rd.,
Flagstaff, June to December, elev. 2150 m, States AHF1/0,
AHF181; Casner Park, October, elev. 2200 m., States
AHF228.
Figure 1. Photomicrographs of Tuber ascospores: (A) Tuber
dryophilum and (B) Tuber levissimum, X520.
402
LITERATURE CITED
Fogel, Robert and J.M. Trappe. 1976. Additions to the
Hypogeous Mycoflora of Colorado. I. Ascomycetes.
Mycotaxon 4:211-217.
Gilkey, H.M. 1954. Tuberales. North American Flora 2, 1:l-
36.
Hall, Joseph G. 1981. A field study of the Kaibab Squirrel
in the Grand Canyon National Park. Suppl. J. Wildlife
Management 45(1):1-54.
Hawker, L.E. 1968. Wall ornamentation of ascospores of
species of Elaphomyces as_ shown by the_- scanning
electronmicroscope. Trans. Brit. Mycol. Soc. 51:493-
498.
Hoffmeister, D.F. and V.E. Diersing. 1978. Review of the
tassel-eared squirrels of the genus Otosciurus. J.
Mammal. 59:403-413.
Keith, J.O. 1965. The Abert squirrel and its dependence on
ponderosa pine. Ecology 46(1 and 2):150-165.
Smith, A.H., Helen V. Smith and Nancy S. Weber. 1981. How
to know the non-gilled mushrooms. 2nd Ed. Wm. C. Brown
Pub.
Stephenson, R.L. 1974. Seasonal food habits of Abert’s
squirrels, Scilurus. aberti. (J, \Atiz. Acad waScr.ae.,
1974.9 Proc eH supp.’.
MYCOTAXON
VOR ROM NO, Ze pope 405-415 January-March 1983
Se eee
STUDIES IN THE GENUS PHOMA. I.
PHOMA AMERICANA SP. NOV.
G. Morgan-Jones and James F. White
Department of Botany, Plant Pathology and Microbiology,
Auburn University Agricultural Experiment Station,
Auburn University, Alabama 36849, U.S.A.
ABSTRACT
Phoma amertcana Morgan-Jones and White is described and illus-
trated from isolates from wheat leaves in Alabama, corn roots in
Georgia and cysts of the nematode Heterodera glyctnes Ichinohe in
Arkansas, Mississippi and Missouri soils.
PROLOGUS
This being the inaugural number of a series of studies in which
taxonomic parameters and species concepts in the genus Phoma Saccardo
are re-examined.
The generic name Phoma has long suffered the ignominious fate of
many other commonly encountered genera of the Deuteromycotina by having
pIde0s volar. arcely indiscriminately, a plethora of “specific epithets
based on such sometimes spurious distinctions as host substrate re-
lationships and miniscule morphological discontinuities. Add to this
the fact that many of the contributions to our knowledge of the genus
have included descriptions ranging from the mediocre to the moderately
competent and we have a recipe for chaos. So much so that a situation
has been reached where the application of specific epithets within the
genus with confidence can, at best, be a hazardous undertaking.
Over the years the main weakness in the documentation of species
peculiarities in the Phoma-complex has been a lack of cultural studies
under standardized conditions and of adequate accounts of phenotypic
plasticity. In Phoma, as in such other genera as Cladosportum Link
and Fusartum Link, it has not been merely a matter of interplay be-
tween "lumpers" and "splitters". There has long been a dearth of
sufficiently adequate information on which to make valid judgements.
With the singular exception of the work of Dennis (1946), critical
comparative studies in vittro of Phoma isolates remained unattempted
Alabama Agricultural Experiment Station Journal Series No. 6-82265
404
until the 1960's when Dr. G. H. Boerema and his co-workers in the
Netherlands initiated their far-reaching reassessment of a chaotic
taxonomy. Dennis, recognizing the impossibility of defining taxa
without linkage with extant type specimens, wisely left in abeyance
the matter of speciation choosing instead to classify his isolated
into groups of strains. During the last two decades Dr. Boerema has
succeeded in doing for Phoma what Dr. W. H. Wollenweber did for
Fusarium some thirty years earlier and we are now in the position
to proceed with evolving a satisfactory classification on 4 sound
foundation.
Much remains to be done to fully document the variable expressions
of taxa given species rank. As more fresh isolates become available
from diverse substrates and ecological niches, in different geographic
locations, opportunity arises to increase the data base by which a
stable, nov to Say practical, taxonomy can come aAnto being,
The investigation by Johnston (1981) on fifteen species of Phoma
occurring on grasses \and legumes in New Zealand pastures is exemplary
of the type of study which should now be undertaken.
In this series we propose describing in detail strains of Prod
oripinatange mainly in the United States.
PHOMA AMERICANA — INTRODUCTION
In the course of studying many isolates from various sources we
have determined seven strains of Phoma, from three diverse subsurare
origins, to represent a taxon for which no name is currently avail-
able iin, thesdiverature?
The first of these strains examined by us was isolated in the
spring of 1981 from leaves of wheat growing in Alabama. The fungus
resembled in some respects the plurivorous Phoma pomorum Thum.
[Phoma pruntcola (Opiz) Wollenw. and Hochapf.] which is cosmopolitan
in its distribution (Morgan-Jones, 1967; Boerema, Dorenbosch and
van Kesteren, 1968, 1971, 1977).
Through consultation with Dr. G. H. Boerema’ we were wiven to
understand that isolates of the same organism had been obtained by
Dr. D. R. Sumner from roots of corn grown in Georgia. subsequently,
during a survey of fungi associated with populations of the cyst
nematode Heterodera glyetnes Ichinohe in Arkansas, Mississippi and
Missouri soybean field soils (Morgan-Jones, Gintis and Rodriguez-—
Kabana, 1981), the fungus was again encountered [it is referred to in
that publication as "black yeast (I)" since, in its cultural char-
acteristics, it somewhat resembles species of the black yeast complex
(sensu de Hoog and Hermanides-Nijhof, 1977) and pycnidia were not
PLATE 1. A, 10-day old colony on MEA; B, 10-day old colony on PDA;
Cc, {-day old colony on cellulose agar; D, clustered pycnidia on celiue
lose agar; E, SEM view of pycnidial cluster; F, droplet-—like deposits
on hypha (stained with aniline blue); G, appendage-like pycnidial hy-
phae; H, uni-ostiolate pycnidia on cellulose agar.
405
406
initially formed].
This Phoma, which is not known to occur in Europe (Boerema, per-
sonal communication), where P. pomorum is quite common, is described
here as a new species.
MATERTALS AND METHODS
All isolates, including three obtained from Dr. D. R. Sumner,
were single-spored prior to use for cultural studies under standard-
ized conditions. Colony characteristics and radial growth rates of
cultures were determined on potato dextrose agar (Difco) and malt
extract agar (Difco) plates incubated at various temperatures (20,
25, and 30 C respectively) in the dark. Four replicates per treat-
ment were used. Cultures were initiated in Petrivdisnes trom ager
dises bearing mycelium, 5mm in diameter (obtained with a No. 2 cork
borer), derived from actively growing margins of 7-day old colonies
growing on PDA. ‘Two cellulose agar [Eggins and Pugh, 1962, but sub-
stituting cellulose (Sigmacel1®) Type 20 for ball-milled Whatman's
cotton cellulose powder] culture plates were prepared for each iso-
late, incubated at 25 C,-and exposed to avcycle or ailvernac iss.
hours near ultra-violet illumination (Westinghouse 20 watt F20T12/
BLB) and 12 hours white light (Westinghouse 20 watt F20T12/CW).
Determinations of radial growth were made after 4 days. Measurements
of pycnidial and conidial size were made from cellulose agar plates
after 10 days.. For each isolate 10 pycnidia and 20 conidia were
measured. Representative cultures (one from each substrate source)
have been deposited at the American Type Cultures Colvecvion.
Pycnidia were embeded in plastic methacrylate. Sections 3 microns
thick were obtained using a Sorvall® JB-4 microtome and stained in
Delafield's hematoxylin followed by congo red. Specimens for scanning
electron micrography were dried using a Denton Vacuum DCP-1 critical
point drying apparatus and coated with gold and palladium using a
Denton DV-502 vacuum evaporator. Micrographs were obtained using a
Phillips AMR model 1000 scanning electron microscope.
TAXONOMIC PART
Phoma amertcana sp. nov.
Maculae amphigenae, ellipticae, brunneae, marginibus rubellis,
2 - 4mm longae. Pycnidia partim immersae in maculis.
Coloniae in agaro maltoso olivaceae, mycelio sparse aereo, post
4 dies 25 C and 19m diametro, reverso olivaceo; in agaro decocto
PLATE 2. Type isolate. A, V.S. bi-ostiolate pycnidium; B, section
of pyenidial wall; C, chlamydospores; D, SEM view ot uniostiolate
pycnidia; E, conidia; F, pycnidial primodium.
407
"
.
-
\
408
tuberorum olivaceae, lanosae vel floccosae, margine sparsae, post
4 dies 25 C ad 53mm diametro, celeriter crescentes, reverso brunneo.
Mycelium ex hyphis septatis, ramosis, subhyalinis vel pallide
brunneis, levibus, 1.5 - 3um crassis compositum; hyphis interdum
irregulariter inspissatis, interdum -fasciculatis. Pyenidia solitaria
vel gregaria vel confluentia, subglobosa vel ampullacea, brunnea
vel fusca, partim inmersa vel superficialia, pseudoparenchymatica,
ostiolata, saepissime collo brevi praedita, 100 - 220um diam.; paries
plerumque cellularum isodiametricarum compositum. Cellulae conidio-
genae (phialides) hyalinae, simplices, ex cellulis interioribus
parietis pycnidii, ampulliformes, 3 - 5 X 3 - 4um. Conidia entero-
blastica, hyalina, simplicia, plerumque biguttulata, cylindrica vel
ellipsoidea, utrinque rotundata, levia, plerumqué continua, interdum
uniseptata, 5-7 X 2- 2.5um. Chlamydosporae intercalares vel
terminales, solitariae vel catenulatae, subglobosae, globosae,
ellipsoideae vel irregulares, uni-, bi- velmulticellulares, cras-—
situnicavae, leves vel asperatac, pallide brunneac vel oiriimnneders .—
HOum diam.
In foliis vivis Tritici, Montgomery County, Alabama, April 1981,
A. K. Hagan, AUA, holotypus.
Pigures! Landes pllaves: | 2 andess
Lesions amphigenous, elliptical, brown, margin reddish in color.
Pycnidia partly immersed in the leaves. Colonies on malt extract
agar (Plate 1, A) olivaceous, with sparse aerial mycelium, particular-
ly in a wide marginal zone, attaining a diameter of 9mm at 20C after
4 days, 19mm at 25 C and 20mm at 30 C, center darker and thinly
floccose or in part slightly ropy in appearance, reverse olivaceciuse;
on potato dextrose agar (Plate 1, B) pale to dark olivaceous, zonated,
lanose or somewhat floccose, margin thin, generally darker towards
the center, attaining a diameter of 39mm at 20 C after 4 days, 53mm
at 25 C and 57mm at 30 C, reverse brownish. On both MEA and PDA
mature pycnidia are produced sparsely after 10 days, partly immersed
in the agar. Colonies on cellulose agar (Plate 1, C) with very little
aerial mycelium, producing abundant pycnidia superficially after 7
days. Mycelium composed of septate, branched, subhyaline to pale
brown, smooth, 1.5 -— 3um wide hyphae; hyphae sometimes irregularly
thickened, occasionally bearing more or less hemispherical or flatten-
ed, droplet-like deposits (Plate 1, F), sometimes aggregated into
closely appressedstrands. Pycnidia solitary or gregarious or fre-
quently, especially on.cellulose agar, confluent, subglobose to flask-
shaped, brown to blackish brown, partly immersed or superficial,
pseudoparenchymatous, covered to varying degrees by hyphae (Plates 1,
E; 2, D; 3, D, E), frequently bearing long, unbranched, flexuous,
appendage-like, 1 - 1.5um wide hyphae which are sometimes slightly
swollen at their tips (Plate 1, G), with one or more ostioles (up to
four), usually bearing one or more short necks, 100 - 220um in
PLATE:.3:. Corn isolate. A, V. S. pycenidiums B; Cy sections of
pyenidial wall; D, E, SEM views of multi - and uniostiolate pycnidia;
F, pyenidial clusters; G, H,. chlamydospores.
iif
/ y
yi?”
410
LD
FIGURE 1. A, V.S.pycnidium of type isolate; B, portion of pycnidial wall
around ostiole (indicated by arrow); C, conidia; D, conidiogenous cells;
E, chlamydospores of type isolate.
411
0
aN
1%
¢,
ye
chlamydospores of
B,
5
1 cluster on cellulose agar
ia
qd
ie
heavily confluent pycnidial cluster.
pyen
C;
3
3
A
solate
FIGURE 2.
i
corn
412
diameter where solitary; where confluent the pycnidial mass (Plates
1, D; 3, F; Fig. 2, A) can reach up to 850um in diameter and contain
as many as 10 to 14 separate units as evidenced by the number of necks;
wall composed of isodiametric or somewhat elongate cells (Plates 2, B;
3, B, C), up to four layers (7 - llym thick), frequently slightly
thicker in the zone of transition between the neck and the main
pycnidial body with individual cells 3.5 - 5um in diameter, wall cells
becoming progressively elongate towards the pycnidial base where
frequently only one layer is present and cells measure 8 -9X1.5 -
2.5um. Conidiogenous cells phialidic, hyaline, simple, smooth-walled,
borne on the innermost cells of the pycnidial wall up to the base of
the neck region (Plate 3, B), subglobose to broadly flask-shaped,
3-5 xX 3 -4um. Conidia enteroblastic, hyaline, simple, frequently
biguttulate, cylindrical or narrowly ellipsoidal, obtuse at cach7end.
smooth, continuous or occasionally one-septate, 5- 7 X 2 - 2.5um.
Exuded conidial mass salmony in color. Chlamydospores very variable,
intercalary or terminal, solitary or in chains, subelobose) ell ipsodial
or irregular, uni-, bi - or multicellular, when septate phragmosporous
or dictyosporous, thick-walled, smooth or roughened, pale brown to
brown, 15 - 4O0um in diameter, non-septate chlamydospores 15 - 18um;
chlamydospores produced abundantly on PDA, sparsely on MEA and
cellulose agar.
On leaves of wheat, roots of corn and in cysts of Heterodera
glyetnes; North America.
Collections examined: on Trttiewn aestivum L., Montgomery County,
Alabama, April 1981, A. K. Hagan, AUA, holotype; on Zea mays L.,
Baker County, Georgia, May 5, 1980, Miller County, Georgia, May 22,
1980, Tift County, Georgia, May 28, 1980, D. R. Sumner, AUA; from
cysts of Heterodera glyetnes from soybean field soils, Lee County,
Arkansas, Pontotoc County, Mississippi, Pemiscot County, Missouri,
July 1981, G. Morgan-Jones and B. O. Gintis, AUA.
ADDITIONAL NOTES
Although the seven strains of Phoma amerteana represent a distinct
taxon, considerable variation is exhibited among them in some regards
particularly in pycnidial and chlamydospore characteristics.
On cellulose agar the strains derived from nematode cysts failed
to show pycnidial clustering and confluence to the same degree as
in the others but multiostiolate pycnidia were abundant. It should
be noted, however, that there is sometimes difficulty in distinguish-
ing between clustered pycnidia derived from the confluence of a few
primodia and bone fide single, but multiostiolate, entities. One
strain from corn differed by failing to produce any pycnidia on this
medium.
The type isolate produced no chlamydospores on cellulose agar,
whereas such structures were produced sparsely on this medium by the
other strains and moderately in the case of the corn isolate that
produced no conidia. The degree of chlamydospore production on
413
MEA and PDA respectively also varies among strains. Furthermore the
type of chlamydospore predominating varies. The corn isolate that
failed to produce pycnidia produced single-celled, heavily melanized,
roughened chlamydospores almost exclusively.
One-septate conidia were encountered, though infrequently, mainly
in ehe cori isolates.
Phoma amerteana is easily distinguished from other species, inclu-
ing P. pomorum, by its chlamydospores, relatively thick-walled
pycnidia, and its slow growth on malt extract agar [P. pomorum attains
a diameter of 5 - 7.5cm on MEA after 7 days (Dorenbosch, 1970) ].
ACKNOWLEDGMENTS
We baveuhnad the benefit. of consultation with Dr. Gerhard! H.
Boerema, Wageningen, The Netherlands, and are grateful to him for his
commemvyeson some of our isolates. Dr. D. R. Sumner, Tifton, Georgia,
kindiveprovided us with cultures of his isolates from Corn Dr,
Richard T. Hanlin, University of Georgia, kindly reviewed the manu-
SCPiDG.
REFERENCES
BOEREMA, G. H., M. M. J. DORENBOSCH AND H. A. VAN KESTEREN. 1968.
Remarks on species of Phoma referred to Peyronellaea. Persoonia
Doieec01—205,
BOEREMA, G. H., M. M. J. DORENBOSCH and H. A. VAN KESTRREN. 1971.
Remarks on species of Phoma referred to Peyronellaea - III.
Persoonia 6: 171-177.
BOEREMA, G. H., M. M. J. DORENBOSCH and H. A. VAN KESTEREN. 1977.
Remarks on species of Phoma referred to Peyronellaea - V. Kew Bull.
31: 533-544,
DE HOOG, G. S. and E. J. HERMANIDES-NIJHOF. 1977. ‘The black yeasts
and allied Hyphomycetes. Stud. Mycol.: 15, Centraalbureau voor
Schimmelcultures.
DENNIS, R. W. G. 1946. Notes on some British fungi ascribed to Phoma
and related genera. ‘Trans. Br. mycol. Soc. 29: 11-42.
DORENBOSCH, M. M. J. 1970. Key to nine ubiquitous soil-borne Phoma-
like fungi. Persoonia 6: 1-14.
EGGING He O: Weand G. J. Fi PUGH, 1962. Isolation of cellulose de-
composing fungi from the soil. Nature, Lond., 193: 94-95.
JOHNSTON, P. R. 1981. Phoma on New Zealand grasses and pasture
legumes. N. Z. Jl. Bot. 19: 173-186.
MORGAN-JONES, G. 1967. Phoma pruntcola. CMI Descr. Path. Fungi Bact.
No. 134. Commonwealth Mycological Institute.
MORGAN-JONES, G., B. OWNLEY GINTIS and R. RODRIGUEZ- KABANA. 1981.
Fungal colonisation of Heterodera glycines cysts in Arkansas,
Florida, Mississippi and Missouri soils. Nematropica 11: 155-163.
MYCOTAXON
WO XN oe NO ice ny Wee eel deat 110 January-March 1983
A NEW SPECIES OF AMANITA
DAVID T. JENKINS
Department of Biology, University of Alabama in Birmingham
Biuningham, AL 35294
ABSTRACT
Amanita media (fig. 1) is described as new from Alabama. It
appears to be intermediate between sections Lepidekla and Vakidae.
Amanita media Jenkins, sp. nov.
Holotype: Alabama - rest area, Hiway 231, between Florala and Opp, Al,
11. vi. 1981, Jeannie and David Jenkins 1629(DTJ).
Corpus fructus medius. Pileus: quanto 78 mm latus, planus con-
vexusque cum in orbe loco depresso tenuique, margo non striatus, pilei-
pellis mediocriter ac facile separabilis, fulgens, levis, argenter ac
alba, verum cum colore destincto vero, praecipue in orbe, caro albue,
quanto 7 mm crassus ad centrum, fastigata adversus marginem. Lamellae:
densae, annexae, gilvae; lamellulae truncae convertentes in superficie,
tenuissime attenuata. Stipes: quanto 50 x 14-15 mm, solidus, gilvus,
levis, tegumen partim et in forma apicis, albus, delicatissimus, iam
evanescens bulbus ad basem obovatus, quanto 30 x 25 mm; reliquae uni-
versi teguminis ita panni fortuiti floccosique in bulbo supero, albae,
facile remotae. Odoratio no distinctus. Spores 4.69-5.47 x (9.37)
10.15-12.50 um.
Fruit body medium, solitary. PILEUS: up to 78 mm diam, plano-
convex with slight depression in center, margin non-striate, pileipellis
fairly easily separable, shiny, smooth, silvery white, but with distinct
tannish tint primarily on disc, flesh white, up to 7 mm thick at center,
tapering toward margin; universal veil as few small, floccose patches.
LAMELLAE: crowded, adnexed, creamy-white; lamellulae truncate to
slightly attenuate. STIPE: up to 50 x 14-15 mm, solid, creamy-white,
smooth; partial veil apical, white, very delicate, soon disappearing;
basal bulb obovate, up to 30 x 25 mm; universal veil remnants as ran-
dom, floccose patches on upper bulb, white, easily removed. Smell not
distinct.
PILEIPELLIS: filamentous hyphae densely interwoven, considerable
gelatinization, 2-6 um diam, moderately abundant, oleiferous hyphae
present. PILEUS TRAMA: filamentous hyphae undifferentiated, moderately
branched, no clamps, 2-7 um diam; inflated cells abundant, mostly ter-
minal, mostly elongate, up to 191 x 38 um. LAMELLA TRAMA: bilateral;
filamentous hyphae undifferentiated, no clamps, 2-8 ym diam; inflated
cells elongate, terminal. SUBHYMENIUM: inflated ramose to subcellular,
no clamps. BASIDIA: up to 43 x 4-11 um, mostly 4-sterigmate, but with
a consipcuous number of 2-sterigmate, thin walled, no clamps. UNIVERSAL
VEIL: filamentous hyphae at base of stipe abundant, sparsely to moder-
ately branched, no clamps, 3-7 um diam, some gelatinization, irregularly
415
LD vere.
® . By
4. i iter
Fig. 1. Amanita media Jenkins (DTJ 1629)
disposed; inflated cells abundant, globose, subglobose, broadly elliptic,
up to 78 x 78 um, very few small elliptical cells, mostly short, ter-
minal chains: tissue on pileus the same. STIPE TRAMA: filamentous
hyphae undifferentiated, sparsely branched, no clamps, 3-6 um diam;
inflated cells terminal, clavate, longitudinally oriented, up to 154 x
4] um, most smaller. PARTIAL VEIL: almost exclusively filamentous
hyphae, moderately branched, no clamps, 2-8 um diam, interwoven;
inflated cells sparse, small, terminal.
SPORES: 4.69-5.47 x ‘(9.37)10.15-12.50 um, (E = 1.99-2.29; Ene
2.12), elongate to cylindric, adaxially flattened, thin walled, hyaline,
amyloid, spore print white; contents guttulate; apiculus sublateral,
cylindrical.
Habitat and distribution: terrestrial, loblolly pine and mixed
hardwoods, Alabama.
Collections examined: rest area, U.S. Hiway 231, between Florala
and Opp, Alabama, 11. vi. 1981, Jeannie and David Jenkins 1629(DTJ):
roadside, U.S. Hiway 231, near Florala, Alabama, 11. vi. 1981, Jeannie
and David Jenkins 1657(DTJ).
Discussion: Amanita media appears to be intermediate between
sections Lepidella and Vakidae. The characters of elongate to cylin-
drical spores, whitish pileus, and a delicate partial veil appear to
indicate a relationship with section Lepidella. However, features such
416
as the easy removal of volval patches from pileus, non-appendiculate
pileus margin, and the pileus margin not exceeding the gills indicates
a relationship with section Vakidae.
Although the above characters are used to delineate sections
Lepidekla and Vakidae respectively, exceptions to the definitiveness
of each character can occur. For exampte, the whitish pileus is most
characteristic of members of section Lepidella. However, Amanita
demissa Corner & Bas, section Validae, has a whitish pileus with a
grayish disc. Singer (1975) mentions that there are a few members of
section Validae with cylindrical spores, a feature more commonly asso-
ciated with section Lepidelka.
The features which characterize section Vakidae, i.e., easy
removal of volval material from the pileus and the non-appendiculate
pileus margin, can occasionally be found in members of section
Lepidekla. For example, the volval remnants of Amanita polypyramis
can easily be washed off by rain or removed by scraping. The appen-
diculate margin is quite definitive when present. The character is the
result of the adherence of the partial veil fragments to the pileus
margin as the fruit body expands. Frequently, however, this material
can easily be washed off or fall off with age, resulting in a non-
appendiculate margin.
It is obvious from the above examples that the variability of
characters within individual specimens is frequent enough that no
single feature can unquestionably indicate a sectional relationship.
As can be seen from the description of Amanita media, it possesses
combinations of characters which indicate a possible relationship with
two different sections. Therefore, Amanita media will not be assigned
to a particular section at this time. Future collections may allow
for better discernment.
ACKNOWLEDGEMENTS
My thanks are extended to Dr. James W. Kimbrough, University of
Florida, for reviewing this article.
EIT ERATURE: CITED
Bas, C. 1969. Morphology and Subdivision of Amanita and a Monograph
on its Section Lepidekla. Persoonia 5(4): 285-579.
Singer, R. 1975. The Agaricales in Modern Taxonomy. 3rd. Ed.
Germany.
MYCOTAXON
VO eA V In NO 2 pp. 417=474 January-March 1983
DESCRIPTIONS OF NEW SPECIES AND COMBI-
NATIONS IN mae fsa AND ERYSIPHE
ZR IEA!
UWE BRAUN
Martin-Luther-Universitdt Halle-Witten-
berg, Sektion Biowissenschaften, WB.
Geobotanik und Botanischer Garten, GDR-
DDR=4020 Halle, Neuwerk 21
The present paper continues the taxonomic studies
in the genera Microsphaera and Erysiphe. It comprises
some descriptions of new species and a new combination.
- Microsphaera neglecta U. Braun spec. nov.
Syn.: M. alni emend. Salmon p.p.; M. penicillata f.
ulmi Jacz. (1927, p.364), nom. nud.; M. ulmi (Jacz.)
Bunkina, in Vodorosli, griby i mhi Dal’nego Vostoka, p.
42, Vladivostok 1978 (not validly published, combination
not indicated, basionym not cited, basionym = nom. nud.).
Mycelium amphigenum, evanescens vel subpersistens,.
Cleistothecia 65-100 pm diam,, cellulae peridii ca. 10-
15 pm diam., appendices 5-12, diametro cleistothecii
1-1.5plo longiores, hyalinae, superne tenui- et inferne
crassitunicatae, basi 5-8 pm latae, ad apicem dichotome
ramosae (4-5x), apice recurvatae, asci 3-6, 40-50 x 30-
40 pm, ascosporae 6-7, 15-19 x 10-12 pm.
Holotypus: hospes — Ulmus americana, U.S.A., Galeria
and Council Hill, I1l., 19-93-1882, Seymour (BPI) ."
Mycelium effused, amphigenous, usually epiphyllous,
evanescent to subpersistent. Cleistothecia 65-100 pm in
diam., cells obscure, polygonal to rounded, ca. 10-15
pm diam., 5-12 + equatorially inserted appendages,
curved and flextous, 1-1.5 times as long as the cleisto-
thecial diameter, stalk hyaline, coloured at the very
base, O-1-septate, moderately thick-walled throughout or
thin above and thick below, 5-8 ym wide near the base,
apex 4-5 x regularly branched, the branchings form a
small, compact complex, tips recurved, primary branches
sometimes somewhat elongated and slightly recured, 3-6
asci, mostly without stalk, 40-50 x 30-40 jm, 6-7 spores,
15-19 x 10-12 pm (fig. 1).
I studied an additional sample of the species (on
418
Ulmus fulva, U.S.A., Columbia, Mo., 9-10-1910, leg. ?,
BPI). The new species resembles M. juglandis-nigrae U.
Braun (distinguished by 4-5-spored ascl). e name M.
enicillata f. ulmi Jacz. has been introduced for the
SEI: on Ulmus americana in N. America. Therefore,
this name is a synonym of M. neglecta. The identity of
Bunkina’s collection from the Far Hast of the Soviet
Union is uncertain.
2. Microsphaera magnifica U. Braun spec. nov.
Syn.: M. alni emend. Salmon p.p.
Mycelium amphigenum, subpersistens. Cleistothecia
(75-) 85-125 wm diam., cellulae peridii 10-25 um diam.,
appendices 5-18, diametro cleistothecii 1-1.5plo longi-
ores, hyalinae, superne tenui- et inferne crassituni-
catae, basi 6-10 pm latae, O-1-septatae, ad apicem di-
chotome ramosae (4-6x), apice recurvatae, asci 4-6, 45-
65 x 40-55 pm, ascosporae 3-5, (16-) 20-26 x 10-15 pm.
Holotypus: hospes - Magnolia acuminata, U.S.A.,
Bureau of Plant Industry, rrisburg, base, Northeast
Erie Co., 18-10-1920, Cubbin (BPI).
Amphigenous, mycelium effused or in spots, subper-
sistent. Cleistothecia scattered, (75-) 85-125 um in
diam., cells conspicuous, 10-25 pm diam., 5-18 equa-
torially inserted appendages, stiff to flexuous, 1-1.5
times as long as the cleistothecial diameter, stalk
hyaline, coloured near the base, smooth, aseptate or
with a single septum at the base, thin-walled above,
thick towards the base, 6-10 ym wide below, sometimes
enlarged at the very base (-15 ym), apex 4-6 x branched,
ornate, somewhat loose, ultimate tips distinctly re-
curved, often somewhat circinate to subhelicoid, 4-6
asci, mostly without stalk, 45-65 x 40-55 pm, 3-5 spores,
(16-) 20-26 x 10-15 pm (fig. 2).
The new species is near to M. nemopanthis Peck, M.
ornata U. Braun and related taxa. It is distinguished
by the features of the asci and ascospores. I investi-
gated two additional samples (on Magnolia acuminata,
U.S.A., Washington, Agric. Grounds, BO-1T-T9TO, Norton,
BPI; on Magnolia liliflora, U.S.A., Semmes, Alab.,
4-11-1946, per).
3. Microsphaera neomexicana U. Braun spec. nov.
Mycelium amphigenum, subpersistens. Cleistothecia
90-140 um diam., cellulae peridii 10-20 ym diam., appen-
dices 10-30, diametro cleistothecii 0.75-1.5 (-25) plo
longiores, hyalinae, tenuitunicatae, eseptatae, basi
5-10 pm latae, ad apicem irregulariter dichotome ramo-
sae (3-6 x), apice non recurvatae, asci 6-16, 45-65 x
25-40 pm, ascosporae 3-6, 18-24 x 10-14 pm.
Holotypus: hospes —- Forestiera neomexicana, U.S.A.,
Plaggstaff, Ariz., 24-9-1919, Godding :
419
Fig. 1-2. Microsphaera neglecta (1), ascocarp, ascus,
two branched Sate of the appendages. Microsphaera
magnifica (2), ascocarp, ascus, appendage. U. Braun del.
420
Amphigenous, mycelium effused, subpersistent. Cleis-
tothecia 90-140 pm in diam., cells of the wall polygonal,
ca. 10-20 pm diam., appendages numerous, about 10-30 per
ascocarp, equatorially inserted or somewhat in the upper
half, radially spread, rather stiff, 0.75-1.25 (-2) x
as long as the cleistothecial diameter, stalk hyaline,
aseptate, rough, thin-walled, slightly thicker towards
the base, about 5-10 ym wide below, apex 3-6 times
branched, branchings very irregular, loose, diffuse,
deeply cleft, tips mostly straight, obtuse, few recurved,
asci 6-16, stalked, 45-65 x 25-40 um, 3-6-spored, 18-24
x 10-14 pm (fig. 33.
The species resembles M. diffusa C. & P. It differs
by relatively short appendages and very numerous asci.
M. neomexicana is also close to M. prasadii Bhatnagar &
Kothari, a species on Hamiltonia in India (distinct by
very numerous appendages).
4. Microsphaera friesii Lév. var. dahurica U. Braun var.
nov.
Syn.: M. alni p.p., ss. Homma (1937); M. friesii
ss. Bunkina (1979).
Cleistothecia 80-105 pm diam., asci 4-5, ascosporae
6-7, 16-24 x 9-12 un.
Holotypus: hospes - Rhamnus dahurica, Japan, Mori-
oka, Prov. Rikuchu, 23-9-1903, Yamada (TNS-F-217398) .1
Amphigenous, mostly epiphyllous, subpersistent.
Cleistothecia 80-105 pm in diam., cells obscure, poly-
gonal to rounded, ca. 8-25 ym diam., appendages + equa-
torially inserted, 7-15, about as long as the cleisto-
thecial diam. (0.75-1.25x), stalk hyaline, sometimes
coloured near the base, smooth, thick-walled towards the
base, thin above, 7-11 pm wide below, apex 4-6 x closely
branched, regular, tips recurved when mature, 4-5 asci,
40-50 x 25-45 um, 6-7-spored, 16-24 x 9-12 ym (fig. 4).
The number of ascospores distinguishes var. dahurica
from var. friesii (3-5). The spores of var. friesii are
smaller.
5. Microsphaera miurae U. Braun spec. nov.
Syn.: M. alni p.p., ss. Homma (1937).
Mycelium amphigenum, subpersistens. Cleistothecia
70-115 pm diam., cellulae peridii 10-20 ym diam., appen-
dices 5-10, diametro cleistothecii 1-2plo longiores,
hyalinae, O-1-septatae, tenuitunicatae, basi 6-10 yum
latae, ad apicem dichotome ramosae (3-5x), apice recur-
vatae, asci 3-5, 40-65 x 30-45 um, ascosporae 4-5, 18-
25exks 12155 yim.
Holotypus: hospes - Lonicera morowii, Japan, Oma-
gari, Prov. Uzen, 3-11-1908, Miura (TNS-F-214110).
A21
Fig. 3-4. Microsphaera
two branched Seata or t
friesii var. dahurica (
. Braun del.
neomexicana (3), ascocarp, ascus,
he appendages. Microsphaera
4), ascocarp, ascus, appendage.
Mycelium amphigenous, subpersistent. Cleistothecia
70-115 pm in diam., cells obscure, ca. 10-20 um dian.,
5-10 equatorial appendages, rather stiff, often deeply
cleft, stalk hyaline, sometimes yellowish, brown near
the base, O-1-septate, thin-walled, slightly thicker
below, 1-2 x diam. of the cleistothecium in length, 6-
10 pm wide below, apex 3-5 x loosely branched, primary
branches usually elongated, often somewhat twisted in
different planes, asci 3-5, without stalk, 40-65 x 30-
45 um, 4-5 spores, 18-25 x 12-15 ym (fig.5).
I examined some additional samples on Lonicera
from Japan (ex TNS): Lonicera morowii, L. orientalis,
L. chamissoi, and L. ciliata. AII collections agree with
WM. miurae. The species iS morphologically related to M.
miyabeana U. Braun on Styrax in Japan and allied taxa.
It 18 not related to M. GEG (cleistothecial appen-
dages with straight ultimate tips) and M. Se
cearum U. Braun (an American species, apex oO e appen-
dages richly branched, compact).
6. Microsphaera ornata U. Braun var. europaea U. Braun
var. nov.
Syn.: (Pseudonym), M. betulae (DC.) Magn., Ber.
deutsch. bot. Ges. 16, p.67 (T1898). ‘
Cleistothecia 75-105 ym diam., appendices 4-13,
diametro cleistothecii 0.5-1 (=1.53plo longiores.
Holotypus: hospes - Betula pubescens, Bavaria,
Bayreuth, Oct. 1874, de Thimen, Ce. unLv. 56° (K).4
Because of the changed "Art. 13" of the ICBN,
adopted at the Sydney Botanical Congress (1981), the
name Microsphaera betulae (DC.) Magn., published as a
combination with the basLtonym Erysiphe betulae DC.
(1815, = Phyllactinia guttata), becomes a synonym of
the latter species. Hence it is necessary to introduce
a new name for the European Betula-Microsphaera. After
seeing a large amount of material on different American
Betula species, I prefer to consider the European fungus
as a mere variety of M. ornata. The collections on Betu-
la lenta and lutea are well distinguished from European
Specimens. Samples on Betula sandbergii and pumila
(studied mat. ex herb. AZ), however, are intermediate.
The cleistothecial appendages on B. sandbergii are
only 0.5-1 times as long as the navctathestas diam.
(1-1.5 x on B. lenta and lutea). The only constant
difference between European and American collections is
in the number of appendages. American specimens possess
always numerous appendages (average >10)."M. betulae"
on B. dahurica in Asia (Bunkina 1979) belongs undoubted-
ly to M. ornata var. ornata.
7. Erysiphe cichoracearum DC. var. poonaensis (Chiddar-
war . braun Stat. nov.
Bas.: Erysiphe poonaensis Chiddarwar, Current Sci.
423
conidia.
ascus,
Erysiphe cicho-
ascus, =
ascocarp,
S.
(5),
endage
ascocarp,
the app
poonaensis,
parts o
raun del
Fig. 5-6. Microsphaera miurae
two branche
racearum var.
424
24(12), p.421 (1955).
Studied mat.: on Goniocaulon glabrum, Poona, India,
IX. 54, P. P. Chiddarwar (IMT 61031, type).
Amphigenous, mycelium persistent, conidia in chains,
ellipsoid to barrel-shaped, 24-29 x 12.5-15 pm (dried!).
Cleistothecia 90-145 um in diam., cells obscure, appen-
dages numerous, basally attached, mycelioid, strongly
interlaced with the mycelium, brown when mature, thin-
walled, septate, simple, ca. 5-10 um wide, shorter than
the cleistothecial diam., numerous asci per fruit body
(ca. 8-12), stalked, 50-70 x 25-35 pm, 2-spored, spores
subglobose, ca. 14-19 x 10-13 um (fig. 6).
Var. poonaensis differs from var. cichoracearum by
small perriie and small, subglobose ascospores. e
differences are only slight; the fungus is hardly more
than a variety of E. cichoracearun.
Literature
Bunkina, I.: Mucnisto-rosjanye griby (sem. Erysiphaceae)
Dal’nego Vostoka. Vladivostok 1979.
Homma, Y.: Hrysiphaceae of Japan. J. Fac. Agric. Hokka-
ido Imp. Univ., 38, 183-461 (1937).
Jaczewski, A.: Karmannyj opredelitel’” gribov. Muénisto-
rosjanye griby. Leningrad 1927.
' I am deeply indebted to the staffs of the mentioned
herbaria (BPI, TNS, K, IMI) that have sent valuable
material for the present investigations.
MYCOTAXON
VOd a Velne INO .bea> Pps 1425-428 January-March 1983
TAXONOMIC NOTES ot ae POWDERY MILDEWS
ital
UWE BRAUN
Martin-Luther-Universitdt Halle-Witten-
berg, Sektion Biowissenschaften, WB.
Geobotanik und Botanischer Garten, GDR-
DDR-4020 Halle, Neuwerk 21
The present paper contains the description of Unci-
nula oleosa Zheng & Chen var. zhengii var. nov. and the
new combination Uncinula necator acta.) Burr. var.
ampelopsidis (Peck) stat. nov. Furthermore, a discussion
abou e impact of the changed "Art. 13" (ICBN) on the
Erysiphaceae is included.
1. Uncinula in Asia on Tilia
Zheng & Chen (1977) described Uncinula oleosa on
Tilia tuan from China. The species is clearly ditrerent
from U. Clintonii Peck, an American fungus on Tilia
americana, by the features of the appendages. v. OLeosa
has few appendages (about 8~23 per ascocarp), whereas
U. clintonii possesses a higher number (average > 20).
e@ appendages of U. clintonii are enlarged from base to
top, including the coiled part. The apical part of the
appendages in U. oleosa is not enlarged, its width is
sometimes even decreasing. Homma (1937) and subsequent
authors determined Japanese collections on Tilia either
as U. clintonii or U. miyabei. I reinvestigated some
specimens on Tilia iakontaa (TNS-F~216134, 216135, 216137).
The material agrees entirely with U. oleosa and is cha-
racterized as follows:
Uncinula oleosa Zheng & Chen, Acta Microbiol. Sinica 17
» pe290 (1977)
Mycelium amphigenous, evanescent to subpersistent.
Cleistothecia 85-125 wm in diam., cells polygonal to
rounded, ca. 10-20 um diam., 8-25 equatorially inserted
appendages, flexuous, sometimes abruptly bent, 1-2.5 x
as long as the cleistothecial diam., mostly 1.5-2 x, hya-
line, coloured at the base, usually 1-septate, thin-walled,
somewhat thicker towards the base, 5-8 um wide below,
width uniform throughout or slightly enlarged upwards
(-10 wm), coiled part not enlarged, sometimes even decre-
asing, apex closely circinate to subhelicoid, oil drops
present, especially in the upper half of the appendages,
426
&
é A
i re ry es
Y OTE ERD
yo aie RY PS
rp . « ©.
a xy J
> és” ‘° of
fey.
ace
ve sae ts aM te
eg BT t6e
Fig.
Vare
ts of
°
>
icziana, two upper par
. U. Braun del.
scocarp, ascus, appendages
ia maximow
B
onica, 4
4. Uncinula oleosa Zheng & Chen var. oleosa on
a
zgnengil on
e appendages
Ti 12a
427
3-7 asci, without or shortly stalked, 45-60 x 35-50 um,
5-7 spores, (15-) 18-25 x 10-15 um (fig. 1).
I studied some collections on Tilia maximowicziana.
The appendages are 5-7.5 pm wide near the base and in-
creasing upwards (- 7-10, rarely - 14 pm). They are
rather regularly increasing from base to top (circinate
part). The apex is frequently slightly enlarged, oil
drops are rare. Otherwise this form of the fungus coin-
cides with typical U. oleosa. It should be regarded as
a variety of the latter species. U. clintonii and U.
oleosa are undoubtedly related, but they are morpho-
logically and geographically differentiated.
Uncinula oleosa Zheng & Chen var. zhengii U. Braun var.
nov. (Etym.: Prof. Zheng, Beijing, Chinese mycologist)
A typo differt appendices ad apicem incrassatae et
clavulatae in forma.
Holotypus: hospes - Tilia maximowicziana, Japan,
Muruyama near Sapporo, 14-10-1925, Homma (TNS-F-216179).
2. Uncinula necator (Schw.) Burr. var. ampelopsidis
(Peck) U. Braun stat. nov.
Bas.: Uncinula ampelopsidis Peck, Trans. Albany
Inet. 7 eipeclomelorc).
Syn.: Uncinula subfusca Berk. & Curt., Grevillea 4,
p.160 (1876).
U. ampelopsidis is usually included in U. necator.
The differences between U. necator on Vitis and the
fungus on Parthenocissus have been pointed out by Sal-
mon (1900) and Blumer (7933). The appendages of U. ne-
cator s. str. are long and flexuous (about 1-6 x as
Tong as the cleistothecial diam.). U. ampelopsidis,
confined to N. America, possesses shorter, rather stiff
appendages (length ca. 1-2.5 x diam. cleist.). The ave-
rage diameter of the ascocarps is larger. The conidial
states are agreeing. There is a close overlapping of
the cleistothecial features in both species, but the
differences are constant. Therefore, I consider U. ampe-
lopsidis as a variety. In Europe (Braun 1982) and Japan
(Nomura 1981) U. necator infects occasionally Ampelopess
or Parthenocissus. Fruit bodies are not formed an e
conidial states correspond with U. necator on Vitis.
3. Nomenclatural notes
The impact of the changed "Art. 13" (ICBN), adopted
at the Sydney Botanical Congress (1981), on the Erysi-
phaceae is only moderate. Two new combinations have been
inevitable. They are already published:
Erysiphe cynoglossi (Wallr.) U. Braun replaces E. asperi-
oliorum Grev.
Sphaerotheca aphanis (Wallr.) U. Braun replaces Sph. al-
428
chemillae (Grev.) Junell
Besides, the citations of some additional species
are influenced. The protected status of names adopted
by Fries is indicated with the sign " : ", It has to be
placed between the original author and Fries. The follo-
wing list comprises the correct citations of the species
in question.
Erysiphe agquilegiae DC. Phyllactinia guttata
up Cichoracearum Dc. (Wallr. : 7.5 Lev.
E. convolvuli Da. Podosphaera clandestina
- depressa (Wallr.) Schlecht. (Wallr. : 7) LeVe
=
bo
Be eopsidis DC. Sphaerotheca epilobii
E. graminis Dc. (= Blumeria (Nalin) Sacc.
graminis C.) Speer) Sph. ferruginea (Schlecht.
x eraclei es ST, une
. hyperici (Wallr.) Blumer S a ss inea (Schlecht.
ae h
(= aera hypericacearum
U. Braun) sph." pacularie ‘nacularis (Wallr. :
E. Les DC,
Be oni. “DC: S ee EROS Seer ety. :
E. ee (Wallr.) Blumer Fr. 5 ter.
Wicrosphaéra astragali (DC.) Unednula. adunca (Wallr.
Trev.
T. Sy... eae
M. berberidis (DC.) Lév. U. bicornis (Wallr.
M@. divaricata (Wallr.) Lév. Tr.) Lev. (= Sawadaea
M. euonymi (DC.) Sacc. bicornis (WalIr. : Fr.)
a rossulariae eee Lév. Homma ) |
Ve fos comc CDC) Wink. U. prunastri (DC.) Sacc.
Ms penicillata (Wallr. : Fr.) 2
2
CV.
I wish to express my gratitude to the staff of the
herbarium TNS for the valuable material that has been
used for the present study.
Literature
Blumer, S.: Die Erysiphaceen Mitteleuropas unter beson-
derer Bee ue ac der Schweiz. Beitr. Krypt.-Fl.
Schweiz 7(1), 1-483 (1933).
Braun, U.: Morphological Studies in the Genus Oidium. III.
Zb1l.. Microbiol. 137, 314-324 (1982).
Homma, Y.: Erysiphaceae of Japan. J. Fac. Agric. Hokkaido
Imp. Univ. 38, 183-461 (1937).
Nomura, Y.: The conidial stage of Uncinula necator occur-
ring on vitaceous plants in Japan. Trans. mycol. Soc.
Japan 22, 333-336 (1981).
Salmon, E.: A monograph of the Erysiphaceae. Mem. Torrey
Bot. Club 9, 1-292 (1900).
Zheng, R.-y. and Chen, G.-q.: Taxonomic Studies on the
Genus Uncinula of China. III. New Species and New
Varieties on Coriariaceae, Euphorbiaceae, Oleaceae,
Sterculiaceae and Tiliaceae. Acta Microbiol. Sinica
17(4), 281-292 (1977).
MYCOTAXON
VOleeWa NO. 2, pp..429-455 January-March 1983
GIGASPORA RETICULATA: A NEWLY DESCRIBED
ENDOMYCORRHIZAL FUNGUS FROM NEW ENGLAND*
R. E. KOSKE
Botany Department, University of Rhode Island,
Kingston, Rl 02651, U.S.A.
DIANE DOUD MILLER
Department of Horticulture, Iowa State University,
mInes-a TAL OUOLS UrS A;
CHRISTOPHER WALKER
21 Barony Street, Edinburgh EH3 6PD, Scotland
Collections of endogonaceous spores from sand dunes
in Rhode Island and an orchard in Massachusetts yielded a
previously unknown species of Gigaspora with unusually or-
nate spores. The species, here named Gigaspora reticulata
sp. nov. forms arbuscular endomycorrhizae.
GIGASPORA RETICULATA Koske, Miller et Walker sp. nov.
(Figs. 1-10)
Sporae cinnamomeae vel obscure castaneae, globosae
vel subglobosae, 188 - 340 x 208 - 470 um, in solo singil-
latim genitae, lateraliter ad cellulam instar suspensoris
efformatae. Sporarum tunica turmis duobus separatis stra-
ta reticulo alveolato inductis. Turma externa stratis tri-
bus. Stratum exterius 0.5-1 um crassum, cinnamomeum vel
castaneum, reticulo 0.5-1 um alto, pagina sporae inter
cristas reticuli spinis gradatim decrescentibus vel cristis
angustis 0.5-2 um altis. Stratum medium 5-11 um crassun,
* Journal Paper No. J-10794 of the Iowa Agriculture and
Home Economics Experiment Station, Ames, lowa.
Project No. 2266.
430
hyalinum vel luteolum, strato interiore 0.3-0.7 um crasso
adhaerens. Turma interna stratis tribus. Stratum exterius
et interius utrumque 1 um crassum, stratum medium 2 um
crassum, amorphum. Cellula instar suspensoris 45-87 x 85-
140 um, cinnamomea vel castanea. Vesiculae in solo 25-30
x 30-40 um, in fasciculis 10-40 numero, atrobrunneae vel
castaneae, projecturis obtusis nodosis. Endomycorrhizae
arbusculis formans.
Spores (azygospores?) orange-brown to dark red-brown,
globose to subglobose, 208-470 x 188-340 um, borne singly
in the soil, laterally on bulbous suspensor-like cells
formed at the tips of thick-walled, sparsely septate or co-
enocytic pale-brown hyphae. Wall structure complex, con-
sisting of two separate groups of wall layers overlain by
an alveolate reticulum. Outer wall group three-layered.
Outer layer 0.5-1 um thick, orange-brown to red-brown,
supporting raised, straight to sinuous interconnecting
ridges that form a reticulum 0.5-1 um high, with 4- to 8-
Sided meshes 2-24 x 2-30 um across. Spore surface between
ridges covered with polyhedral, conical or subcylindrical
spines, or narrow straight, curved, or angular ridges 0.5-2
um high and 0.25-0.5(-1) um apart; the spines 0.3-1.5 um
diam at base, tapering to a point or a rounded tip less
ORE TE RSE LES GRO BESTE LG SSE TE ES BEEPS EL SRI BEER TTY ES ERE EADIE ESE I LORI EET EI, GEOG BT ELIS LE LIE ELE LET IS LEE AS PELE, SEE LEIS
FIGURES 1-9. Gtgaspora reticulata spores by brightfield (B), differential
interference contrast (DIC), or scanning electron (SEM) microscopy.
i An intact spore showing the reticulate outer covering and the laterally
attached bulbous suspensor-like cell (B).
Be The vesicles formed in the soil by G. reticulata.
Se A crushed spore that has lost or failed to develop fully the reticulate
outer covering, the vestiges of which (ret) can be seen near the spore
base. The flexible inner wall-group (iw) and the germination shield (gs)
can clearly be seen (DIC).
4. A crushed spore showing the reticulate covering and the flexible inner
wall group (iw) (B).
5. Detail of the outer wall-group. The alveolate reticulum, spiny layer,
and thick, colored middle layer are evident. The thin, hyaline basal layer
is not evident (SEM).
6. Base of a spore showing the bulbous suspensor-like cell with a peg-like
hyphal protrusion (SEM).
7. Spore surface detail by brightfield microscopy.
Si Same part of spore surface as in Figure 7, but by differential inter-
ference microscopy.
9. Detail of reticulum showing alveoli filled with spines.
431
432
FIGURES 10-12. Light microscopy of outer wall-layers of Gigaspora reticulata
(Figure 10) and G. nigra (Figures 11 and 12). Figures 11 and 12 show the
even, rounded holes in the outer wall of the latter. Figure 10 shows the
difference between the reticulate outer covering of the former and the
apparently sinuous nature of the wall of G. nigra beneath the outer wall.
can be compared with the spines and ridges evident on G. rettculata in
Figure 10.
This
433
than 0.5 um diam; the ridges 0.5-1 x 1-7 um at the base,
tapering to a rounded edge less than 0.5 um wide. Middle
layer hyaline to pale yellow, 5-11 um thick, tightly adher-
ent to the thin, hyaline inner layer that is 0.3-0.7 um
thick. Reticulate ridges on outer wall supporting a de-
tachable alveolate reticulum 0.5-2 um wide and 2-6 um high.
Inner wall group three-layered, consisting of membranous
inner and outer hyaline layers each 1 um thick, connected
by a hyaline amorphous middle layer 2 um thick. Outer and
inner layers separating before germination to form a com-
plex, circular to reniform germination shield, 94-156 x
156-208 um, from which germ tubes arise near the spore base.
Suspensor—-like cell 45-87 x 84-140 um, with a peg-like
protrusion extending 10-20 um towards the spore wall. Wall
of suspensor-like cell laminated, orange-brown to red-brown,
3-5 um thick except near the spore-base where it thickens
to 7-10 um; in some specimens, consisting of two distinct
laminated layers, the inner layer 1.5-3 um thick, orange-
brown to red-brown, the outer layer 2-7 um thick, hyaline
to pale yellow.
Vesicles in the soil forming in clusters of 10-40 on
somatic hyphae or around the suspensor-like cell, dark brown
to red-brown, 25-30 x 30-40 um, with blunt, knobby projec-
tions 4-8 x 2-6 um.
Forming endomycorrhizae with arbuscules.
DISTRIBUTION AND HABITAT: Known from coastal sand
dunes at the boundary with a saltmarsh, Sand Hill Cove,
Rhode Island, and from a barrier sand dune at Moonstone
Beach, Rhode Island. Also found in a Montauk soil (coarse-
loamy, mixed mesic Typic Fragiochrepts (Soil Survey Staff,
1975) ) in an orchard at Belchertown, Massachusetts.
Associated, but not proved mycorrhizal, with roots of Phrag-
mites communis Trinx, Ammophila breviligulata Fern., Myrica
pensylvanic Loisel., Malus domestica Borkh., Prunus
persica (L.) Batsch., Dactylis glomerata L., and Agropyron
repens (L.) Beauv.
MYCORRHIZAL ASSOCIATIONS: Forming arbuscular mycor-
rhizae in greenhouse pot cultures with Sorghum sudanese
(Piper) Staph. and Coleus x hybridus. Attempts to induce
mycorrhizal formation with Allium sativa L. cv Agway sweet
spanish, Zea mays L. cv seneca chief, and Malus domestica
were not successful.
434
ETYMOLOGY: Latin, reticulata, referring to the prom-
inent reticulate outer wallcovering of the spores.
COLLECTIONS EXAMINED: HOLOTYPE: RHODE ISLAND -
Washington County, Sand Hill Cove, in rhizosphere of Phra-
gmites communis on a sand dune. Collected 23 iii 1981
(Koske Collection # 286, Walker Accession # 379) (OSC; iso-
types FH, K). PARATYPE: MASSACHUSETTS - Hampshire County,
Belchertown, University of Massachusetts Horticulture Re-
search Farm. Collected 13 viii 1980 (Miller accession # 56,
Walker # 321) (OSC, ISC). OTHER COLLECTIONS: RHODE ISLAND,
Washington County, Sand Hill Cove sand dunes, 4 iv 1979
collected by E. Ovsenik (Koske # 239): RHODE ISLAND -
Washington County, Moonstone Beach sand dune, 9 ix 1978
(Koske # 201). In addition, specimens from pot cultures
established with sorghum and coleus have been examined. Be-
sides the specimens lodged in herbaria, samples have been
retained in the collections of each suthor.
Gigaspora reticulata is readily distinguishable from
other species in the genus by the prominent dark reticulum
covering the spore surface. Only G. nigra J. F. Redhead
in Nicolson & Schenck could be confused with G. reticulata,
but. these two differ’distinctly. CG. reticulata nastelos
bose to subglobose spores 188-340 x 208-470 (mean 264 x
281) um, considerably smaller than spores of G. nigra,
which are globose and 297-1050 (mean 402) um in diameter.
In addition, spores are much darker than those of G. reti-
culata , being dark brown to black compared with orange-
brown to dark red-brown. The main difference between the
species, however, is in the wall structure. G. reticulata
has an outer wall covered by rounded spines and ridges that
are contained, but not overlain, by a polygonal reticulum
with ridges 0.5-2 um wide and meshes varying from 2 x 2 to
24 x 30 um across (Figure 10). The inner wall-group of G.
reticulata consists of two membranous walls apparently
fastened together by an amorphous middle layer. G. nigra,
in contrast, has a distinct outer wall pitted by rounded
pores 7-10 um diam, overlaying a second wall made up of
coiled elements (Nicolson & Schenck, 1979; Old et al., 1973)
that appear as densely packed, sinuous rods by light micro-
scopy (Figures 11 and 12). The inner wall-group in this
species consists of a continuous, laminated wall surround-
ing two thin, membranous walls.
435
G. reticulata spores tend to lose their suspensor-like
cells during sieving from the soil and can then be confused
with spores of fungi in the genus Acaulospora. The resem-
blance of this species to A. bireticulata Rothwell &
Trappe is particularly striking, and the plates in Rothwell
and Trappe (1979) and Walker and Trappe (1981) (Figure 2
amd Figure 12, respectively) are almost indistinguishable
from Figures 7, 8, and 10 in this paper. Care should there-
fore be taken when identifying spores in this group to
verify the nature of the attached hyphae.
ACKNOWLEDGMENTS: We thank Dr. William J. Lord, Extension
Pomologist, University of Massachusetts, who kindly allowed
the collection of samples at the Massachusetts Horticul-
tural Research Farm; E. Ovsenik for collecting one of the
sand samples from Rhode Island; and Don Scales for the
scanning electron microscopy. We also thank Dr. N. C.
Schenck and Dr. J. M. Trappe for their helpful comments.
Dr. Trappe prepared the Latin diagnosis. Dr. Francis
Sanders suggested the term "germination shield" and kindly
permitted its use in this description.
LITERATURE CITED:
Pieter. vi. uNtCcOLson, andsJ . F.) Redhead GEG 73)! oo uh
new species of mycorrhizal Endogone from Nigeria with
a distinctive spore wall. New Phytologist 72: 817-823.
Nicolson, T. H. and N. C. Schenck (1979). Endogonaceous
mycorrhizal endophytes in Florida. Mycologia 71: 178-
193.
Rothwell, F. M. and J. M. Trappe (1979). Acaulospora
bireticulata sp. nov. Mycotaxon 8: 471-475.
Soil Survey Staff (1975). Soil Taxonomy. USDA Soil Con-
servation Service Handbook No. 436.
Walker, C. and J. M. Trappe (1981). Acaulospora spinosa
sp. nov. with a key to the species of Acaulospora.
Mycotaxon 12: 515-521.
MYCOTAXON
Vol. XVI, No. 2, pp. 436-440 January-March 1983
A NEW THERMOPHILIC SPECIES OF MYCELIOPHTHORA
TAKEYOSHI AWAO? AND SHUN-ICHI UDAGAWA
1 Central Research Laboratories, Ajinomoto Co.) ince
Suzuki-cho, Kawasaki-ku, Kawasaki-shi 210, Japan
2 National Institute of Hygienic Sciences, Kamiyoga
l-chome, Setagaya-ku, Tokyo 158, Japan
SUMMARY
A new hyphomycete, Myceliophthora hinnulea Awao &
Udagawa isolated from cultivated soil in Japan, is describ-
ed. The species is typically thermophilic, with a minimum
growth temperature somewhat below 20 C, an optimum between
40 and 45 C, and a maximum somewhat above 50 C.
In the course of a study of thermophilic fungi in
Japan (Awao and Mitsugi, 1973; Awao and Otsuka, 1973,
1974), a hyphomycete belonging to the genus Myceliophthora
was isolated from cultivated soil. It differs from previ-
ously described species by having dull red to grayish brown
colonies and brownish conidia with a conspicuously verru-
cose to spinulose surface.
Myceliophthora hinnulea Awao §& Udagawa, sp. nov. (Fig. 1)
Coloniae in agaro cum decocto tuberorum ad 40 C
effusae, paene planae, constantes ex mycelio basali tenui
et conidio abundanti, velutinae vel plus minusve floccosae,
primo albae vel roseo-vinaceae, dein griseo-brunneae vel
hinnuleae, ad marginem tenues et late submersae; reversum
brunneum vel hinnuleum vel griseo-sepiaceum.
Mycelium ex hyphis hyalinis, ramosis, laevibus, septa-
tis, 1.0-5.0 (-6.5) um diam compositum; hyphae fertiles
plerumque aeriae, dense ramosae, 1.5-3.0 wm diam. Cellulae
conidiogenae in hyphis gerentes, terminales vel laterales,
sessiles vel saepe brevipedicellatae, ad apicemisaepe
inflatae usque 4.0-6.0 x 3.0-4.0 um et ampulliformes.
Blastoconidia plerumque 1-4 per cellulam conidiogenamn,
solitaria vel in 2-3 catenata, primo hyalina vel dilute
437
flava et laevia, dein flavo-brunnea vel brunnea, incras-
sata, subglobosa vel ovata vel pyriformia, (7.0-)8.0-9.0
(-12.0) x (5.0-)6.0-7.5(-10.0) um, ad basim cum contractis
Cicatricibus, irregulariter et, conspicue verrucosa vel spi-
nulosa; spinae usque 1.0-4.0 um longae.
Holotypus No. AJ-6773, isolatus e solo sativo, Nira-
yama, Shizuoka, in Japonia, in 24.11.1973, leg. T. Awao.
In@iercb< NHL,
Etymology: lat. hinnuleus = fawn, referring to the
colony color.
Colonies on potato-dextrose agar growing very rapidly,
attaining 7.8-8.0 cm diam within 3 days at 40 C and more
than 8.5 cm diam within 10 days, almost plane, consisting
of a thin basal felt of vegetative mycelium and abundant
conidia-bearing structures, velvety to more or less floc-
COcc el ooscly covered with rather sterile aerial hyphae, at
first white to’dull red: such as Aurora (M. 10B4; ct.
Kornerup and Wanscher, 1978) or Rosy Vinaceous (Rayner,
1970), becoming grayish brown (M. 7D3) or Fawn (Rayner,
1970) in age; margin thin and broadly submerged; exudate
limited, clear to pale yellow; reverse brown (M. 6E4) or
Fawn to Grayish Sepia (Rayner, 1970).
Mycelium composed of hyaline, branched, smooth-walled,
Septave es! 0->.0(-6.5) um diam hyphae; fertile hyphae
mostly aerial, often densely branched, 1.5-3.0 um diam.
Racquet hyphae absent. Conidiogenous cells borne terminal-
ly or laterally as side branches on hyphae, often with
short or long pedicels which are terminally inflated into
ampulliform swellings measuring 4.0-6.0 x 3.0-4.0 um.
Blastoconidia usually 1-4 borne on ampulliform swellings,
occasionally on short protrusions, solitary or in chains of
2-3, at first hyaline to pale yellow and smooth-walled,
becoming yellowish brown to brown, thick-walled, subglobose
to ovate or pyriform, (7.0-)8.0-9.0(-12.0) x (5.0-)6.0-7.5
(-10.0) um, with narrow basal scars, irregularly and con-
Sspicuously verrucose to spinulose; spines up to 1.0-4.0 um
in length.
Colonies on YpSs agar growing very rapidly, attaining
7.5-7.7 cm diam within 3 days at 40 C and more than 8.5 cm
diam within 10 days, with surface appearing more floccose,
producing abundant conidia on the basal felt and aerial
hyphae, at first white to reddish gray (M. 8B2) or Pale
Vinaceous (Rayner, 1970), later becoming dull red to gray-
ish brown (M. 8D3 to M. 9C4) or Vinaceous Buff (Rayner,
1970); exudate more abundantly, mostly as small droplets
adherent to the mycelium, clear to pale yellow; reverse
brown (M. 6ES) or Umber (Rayner, 1970).
.
Hy
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:
a
ia of Myceliophthora
Conidiogenous cells and conid
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-6773 strain
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439
Cellulolytic, but not distinctly keratinolytic.
Thermophilic with optimal growth at 40-45 C, minimum
somewhat below 20 C, maximum somewhat above 50 C (Table 1).
Table 1. Radial growth and conidia formation of Mycelio-
phthora hinnulea strain AJ-6773 at various temperatures
Temper- Growth, PDA Conidia,PDA Growth,YpSs Conidia,
ature YpSs
(C) 3d 10d 3d 10d Sd i tOda Ge Schram Od
17 0 0 0 0
20 0 5 ty 0 5 oo
ZO 10-12 65-67 - + 12-14 65-70 i +
30 25-26 >85 + ++ 28-29 >85 + os
on 61-63 >85 ++ +++ 59-60 >85 ++ oar
40 78-80 >85 +++ ++++ 75-77 >85 ++ +++
45 80-85 >85 +4+4+4+ ++4++ 72-75 >85 cae +++
50 48-50 >85 + ++ 42-45 >85 a ++
55 0 0 0 0
Growth: colony diam (mm) after 3 and 10 days.
Conidia: ++++, very abundantly produced; +++, abundant;
Peeoederacc,. + tslight; +, very slight; and =) not
produced.
Isolation: cultivated soil, Nirayama-machi, Tagata-
Sun wonszuoka-pref., Japan, 24 Feb. 1973, coll. T. Awao,
No. AJ-6773 (holotype). The specimen studied is preserved
at the Mycological Herbarium, National Institute of Hygie-
nic Sciences, Tokyo (NHL). Subcultures of AJ-6773 have
been deposited with the American Type Culture Collection
(ATCC) and the Centraalbureau voor Schimmelcultures (CBS).
The genus Myceliophthora was reintroduced by van
Oorschot (1977) to accommodate three Chrysosporium-1like
hyphomycetes possessing blastoconidia with narrow basal
scars and often with the remains of ampulliform swellings
produced as a supporting cell. Among these, two have been
known as the anamorphs of two thermophilic Ascomycetes, viz.
Thielavia heterothallica Klopotek and Corynascus thermophi-
lus (Fergus §& Sinden) Klopotek. She has, more recently in
her taxonomic revision of Chrysosporium and allied genera
(van Oorschot, 1980), added five species and provided a key
440
to species currently accepted in the genus. Almost all
hitherto known species are more or less thermotolerant or
thermophilic, with sporulation often being good between 30
and 40 C.
Myceliophthora mostly closely resembles M. vellerea
(Sacc. §& Speg.) van Oorschot and M. thermophila (Apinis)
van Oorschot, the anamorph of Thielavia heterothallica.
It differs from the former in the rich ‘pinkish color of
its young colonies and the rapid growth at high tempera-
ture, because the growth temperature of M. vellerea ranged
between 10 and 30 C. On the other hand, maximum growth of
M. thermophila occurs at temperature of 50 C (our examined
strains of M. thermophila: CBS 117.65 and UAMH 2015 =
semeniuk No. ° 200-2)"; In this respect, ‘the latter is, muca
Similar to the new species but there are some discrepan-
cies in the colonies color and the conidial morphology
such as shape and ornamentation. The conidia of M. ther-
mophila differs also from those of M. hinnulea by the
hyaline color. Finally, because of the conspicuously ver-
rucose-spinulose character of conidia, M. hinnulea is dis-
tinct from the another thermophilic species, M. fergusii
(Klopotek) van Oorschot, the anamorph of Corynascus thermo-
philus, and the other species having smooth conidia.
ACKNOWLEDGMENT
We would like to thank Professor Michael R. Tansey of
the Indiana University for critically reviewing the manu-
Script,
LITERATURE CITED
Awaog 1.3 and) Ke Mitsugi. 1973." Notes=on thermopniuiie
fungi in Japan (1). Trans. mycol. Soc. Japan 14: 145-160.
Awao, T., and S. Otsuka. 1973. Notes on thermophilic
fungi in Japan (2). Trans. mycol. Soc. Japan 14: 221-236.
Awao, T., and S. Otsuka. 1974. Notes on thermophilic
fungi in Japan (3). Trans. mycol., Soc. Japan’ [5ag/-27-
Kornerup, A., and J. H. Wanscher. 1978. Methuen Handbook
of Colour. 3rd ed. E. Methuen, London.
Oorschot, G.- A. N; van.) 1977. The, genus. Myceliophthoz an
Persoonia 9: 401-408.
Oorschot, C. A. N. van. 1980. A revision of Chrysosporium
and allied genera. Stud. Mycol. 20: 1-89.
Rayner, R. W. 1970. A Mycological Colour Chart. Common-
wealth Mycological Institute, Kew, Surrey § British Myco-
Lovicalesociety .
MYCOTAXON
Vol. XVI, No. 2, pp. 441-460 January-March 1983
OPERCULATE DISCOMYCETES FROM RANA (NORWAY) 5.
RHODOSCYPHA GEN. NOV. AND RHODOTARZETTA GEN. NOV.
HENRY DISSING
Instttut for Sporeplanter, K¢ébenhavns Untversttet
0. Fartmagsgade 2D, DK-1353 Kébenhavn K., Denmark
and
SIGMUND SIVERTSEN
Det Kongeltge Norske Videnskabers Selskab, Museet,
Botantsk Avdeling, Untversittetet t Trondhetm,
N-7000 Trondhetm, Norway
ABSTRACT
Rhodoscypha gen. nov. and Rhodotarzetta gen. nov. are
described; Rhodosecypha ovtlla (Peck) comb. nov., Rho-
dotarzetta rosea (Rea) comb. nov., and Weotttella
aphanodtetyon (Kobayasi) comb. nov. are proposed. De-
limitation of the genera Leucoscypha, Neotttella and
Rhodoscypha is briefly discussed.
INTRODUCTION
During the 6th European Mycological Congress 1974 in Avig-
non, France the idea was presented that Pegztza ovtlla Peck
is identical with Hwnarta rhodoleuca Bres. (Dissing 1975).
Brome the beginning of our joint studies of the flora of
Pezizales in the municipality of Rana, Norway (cf. Dissing
& Sivertsen 1975) we have considered it necessary to pro-
pose a new genus for this taxon. A formal proposal of the
new genus Rhodoseypha has until now been postponed because
we wanted definitive results from cytochemical analyses of
carotinoid pigments that clearly supported a separation of
Rhodoseypha ovtlla and species of the genus Neotttella
(Cooke) Sacc.
442
Such evidence is hardly obtained by the preliminary results
presented here, but new descriptions and proposals are the
result of our desire to incorporate these new taxa in a
proper way in the coming flora on "Nordic Macromycetes,
order Pezizales".
MATERIALS AND METHODS
Fresh material of Rhodoscypha ovtlla, Rhodotarzetta rosea,
and Neotttella aphanoditctyon has been studied. Dried mate-
rial has been supplied by the following museums and her-
baria: Botanical Museum, Copenhagen (C), Plant Pathology
Herbarium, Cornell University, Ithaca (CUP), Rijksher-—
barium, Leiden (L), Herbarium, The New York Botanical
Garden, New York \(NY), Herbarium, New York State Museum,
Albany (NYS), Botanical Museum, Oslo (0), Department of
Botany, Panjab University, Chandigarh (PAN), Swedish Museum
of Natural History, Stockholm (S), Institute of Zoology and
Botany, Tartu (TTA), Museum of the Royal Norwegian Society
for Science and Letters, Trondheim (TRH), and The Her-
barium, Institute of Systematic Botany, University of
Uppsala (UPS). Madam F. Candoussau, Pau, France, generously
supplied material from her private herbarium.
Treatment of the material is in accordance with Dissing &
Sivertsen (1980:101).
Rhodosecypha Dissing & Sivertsen gen. nov.
Carposoma dtsetforme vel cupulatum margitne promtnente, ses-
stle vel substtpttatum, extra ptlits flexuosts membrants
distromaticts vestittum; hymentum roseolum, raro albtdum.
Exetpulum onmino textura tntricata, extra ptlts obtusis
saeptatts ramtficatts vestttum membrants tntertortbus
tenutbus aequaltbus, extertortbus ecrassts tnaequaltbus
tineturam Cotton Blue et t. carmineam decltnanttbus.
Aset operculatt, non amylotdes, maturt non prominentes.
Paraphyses saeptatae, supra ramtftcatae, tnfra anastomosan-
tes, cellula cugusque aptcalt guttulas extguas numerosas
continente.
Sporae untsertatae, magnae, fustformes, laeves vel verru-
ets subttltbus solttartts ornatae, quaeque guttulas primum
duas magnas, matura numerosas minores conttnens.
443
Figure 1. Rhodoscypha ovilla.
PruLienOog1es,. MO. / 5,01) (TRH) Sarl;
Nuclet omnes titneturan carmtnean captantes, tn sports
UNC, alt DL Ssinguli — 7nt.
Habttat tn humo vel frustults plantarum, raro tn ltgno.
Spectes typtftca Rhodoscypha ovtlla (Peck) Dissing & Si-
vertsen comb. nov., bastonyma Peztza ovtlla Peck 1875
GLB 7b). 66.
Fruit bodies disc-shaped or cup-shaped, with prominent mar-
in: hymenium pinkish, rarely whitish, outside with
flexuouse hairs with two-layered walls. Sessile or sub-
stipitate.
Excipulum throughout of textura intricata; outside with
septate, branching, blunt hairs with a thin, regular inner
Wadleand ay whitish, ‘thick; irregular, cyanophobic. and: car-
minophobic outer wall.
444
Asci operculate, non amyloid, not protruding when mature.
Paraphyses septate, branching above, anastomosing below, in
the uppermost cell with numerous small guttules.
Spores uniseriate, large, fusiform, smooth or delicately
ornamented with isolated warts, at first with two large
guttules, when mature with numerous smaller ones.
All cells with carminophilic nuclei; spores uninucleate,
other cells with one to seven nuclei.
On soil-or plant. debris, rarely on wood:
Type species Pezgtza ovilla Peck.
DISCUSSION
Harmaja (1977) included Peziza ovilla in the genus Leuco-
seypha Boud. emend. Rifai (1968). Svrcek (1974) did the
same by including the synonymous species Humarta rhodoleuca
Bres. Harmaja correctly argued that cytologically, t.e. the
carminophilous nuclei of the spores and other cells and in
excipular anatomy Leucoscypha leucotrtcha (Alb. & Schw. ex
Fr.) Boud., the type species of Leucoscypha, and P. ovtlla
are much alike.
On the other hand we find the two species differing signi-
ficantly in a number of other characters, viz. the hairs,
paraphyses, spores, and habitat, in a way which justifies
the proposal of the above new genus Rhodoscypha for Peztza
OUT LLG.
In L. leucotrtcha the hairs are stiff, with = twomve. =
defined regular layers in the wall, in P.-ovt/lothesones
layer is thin and regular, while the outer layer is unique
among the Pezizales in being extremely thick, irregular, of
an amorphic, unstainable substance. The paraphyses in P.,
ovtlla often have the uppermost cells densely filled with
numerous refractive, pale reddish brown guttules. In lL.
leucotricha there are a few, colourless guttules in “the
uppermost cells. In Leucoscypha species the spores are per-
manently uni- or biguttulate, in P. ovtila they are at
first biguttulate, then multiguttulate.
445
Hi
‘i wonigganill
doscypha ovtlla.
MO 73
Rho
Cotton Blue.
Figure 2
1 section
x 200
)
e
(
ThE
iki)
ina
Marg
446
Finally P. ovtlla may be characterized as a calciphilous,
montaneus (Svrcek 1974) and subarctic species, while JL.
leucotrtcha mainly is confined to acid soil in temperate
‘areas.
Rifai (1968) and Eckblad (1968) independently circumscribed
the genus Leucoscypha in a way which included species of
the genus WNeotttella (Cooke) Sacc. Species from these
genera have the same excipular anatomy, whitish hairs on
the outside, and carminophilous nuclei in spores and other
cells. On the other hand they differ significantly in at
least a couple of other features, viz. cytochemistry and
biology.
Arpin (1969) showed\Neotttella rutilans (Fr.) Dennis’ ‘to
possess B- and y-carotenes as the quantitatively most
important pigments. To our knowledge no similar analyses
have been performed for L. leucotrtcha, but it appears un-
likely that the above pigments, or others characteristic
for N. ruttlans (Arpin, 1.c.:104) would be demonstrated in
L. tLeucotrtcha. As pointed out. above’ UL. jleucotrichia mis
confined to acid substrates, ‘often on bare *sorl, whe tenn.
ruttlans and its allies always are associated with mosses,
mostly species of Polytrtchum.
It should be noted that in our preliminary analyses of
pigments in R. ovtlla (Hammerness, MO 79.60) made by thin-
layer-chromatography and spectrophotometry, it was possible
to demonstrate four different, unidentified, pigments.
Quantitatively most important was a winered to carmin
pigment with an absorbtion maximum at 490 nm. It was not
possible to demonstrate B- and yY-carotenes. W. ruttlans
served as reference material.
In spite of the mentioned similarities between taxa treated
by others as Leucoscypha species we prefer to consider the
smaller, more natural genera Leucoscypha Boud., Neotttella
(Cooke) Sacc. and Rhodosecypha Dissing & Sivertsen.
Figure 3. Rhodoscypha ovtlla.
Nuclei in carminoaceticacid. a. Spores. b. Paraphyses.
Serato MOS 7s: (Cr aqo,. 1085.
Rhodoscypha ovilla (Peck) comb. nov.
Pezizq, tovilia Peck,. Anns Rep. (N.Y. (State Mus. 28:66,
PSS eel G1 Gt
Weeurrella Ov1LLa (Peck) Sace.; Syll. Fung. 8:194, 1889,
Patella ovtlla (Peck) Seaver, The North American Cup-fungi
(Operculates) :163, 1928.
Leucoseypha ovilla (Peck) Harmaja, Karstenia 17:73, 1977.
mumonitas=rnodoteued ‘Bres:,;. <Fungi Trid« ©11:79, \ 2892.
Pustularta rhodoleuca (Bres.) Boud., Hist. Class. Discom.
mena. 535/907:
Leucoseypha rhodoleuca (Bres.) svrtek, Ceska Mykol. 28:132,
1974.
Figures 1-5 & 7.
448
Fruit bodies.0.5=L cm broad, solitary or /inesmall clusters,
at .finst globose: “with an.'irregular, “siic-like sopenina,
later expanding, disc-shaped to cup-shaped, hymenium pin-
kish, rarely whitish, margin prominent, crenulate, irregu-
larly dentate, whitish, outside covered by a dense mat of
short, whitish hairs; sessile or substipitate.
Outer excipulum of textura intricata, individual cells
10-20 um broad, interwoven, content staining blue in Cotton
Blue; towards the margin, the cells tend to form subparal-
lel rows with the outermost cell club-shaped.
Medullary excipulum as the outer excipulum, cells densely
interwoven, 10-15 um broad, content staining weakly blue in
Cotton Blue. There is no sharp delimitation between outer
and inner excipulum.
Hairs ~2}50=230 wim long; -15-17 Um broad,, flexucts 7 opm
with two-layered walls; outermost wall 5-6 um thick,
whitish, continuous, surface irregular, rough; innermost
wall’ thin; regular; content. of cells staining bluesinagcou.
ton Blue and yellowish in Melzer's reagents; the outermost
wall do not stain in any of the reagents; hairs with few
septa, branching and sometimes anastomosing, each cell with
1-7 carminophilous nuclei. Subhymenium 60-85 um broad, of
densely interwoven cells, some of which are staining heavi-
ly blue in Cotton Blue.
Hymenium 385-425 um high; asci 14-17 wm broad, cylindric,
thick-walled, non-amyloid, the operculum being circular,
5-6 um broad; asci not protruding when mature.
Paraphyses septate, above branching, slightly enlarged to
4-5.3 um broad, below anastomosing; in fresh specimens the
uppermost cell has numerous small pale reddish brown gut-
tules; each cell in the paraphyses has 2-4 (rarely 1 or
5-6) carminophilic, subglobose to ellipsoid, nuclei, 2-4 X
5-7 um.
Spores) uniseriate, (33-39...7-4632° < lO=L34-16e5 imeeersee
form, much varying in size and shape, with one, 6-8 Um
broad, carminophilous nucleus; spores at first with two
guttules, when mature with many small guttules, which in
dried, rehydrated material merge into one large, central
guttule. In the light microscope the spores appear smooth,
but in the scanning electron microscope they prove to be
very delicately ornamented with low isolated warts.
Figure 4. Rhodoscypha ovtlla.
a. Mature spores from fresh material in H20. MO 75.61
(TRH). b. Hairs with thin inner wall (arrows) and
Eiyeckn Outer wall, ~<ni 2s KOHS. MO 7232.73) (C)i. <c.. (Para=
physes from fresh material, with numerous’ refractive
Cube les. MO 75.601 (TRA). ds Spores, anrCotton. Blue;
fromptype of Pe2zt2q ovLLia (NYS)@ e@. Spores, in Cot-
ton Blue, from type of Humarta rhodoleuca, (S). a &
Peron by ane ee, x. 1085.
MATERIAL
The above description is based mainly on the following col-
lections which have been studied in detail:
Norway: Nordland, Rana, Store Alteren, map sheet 1927 I, VP
Done cepcember) 1972, MO. 72.137 (TRH); -~oabid., 5S .septem=
Bema oo, MOUS. 73(C)'s.— ibid... 8 September, 197577 MO 75.61
(C,CUP,H,NY,NYS,O,S,UPS,TRH,ZT); - @rtfjellmoen, map sheet
2027" 1V, VP 86,\9 September 1975, MO 75.68 (TRH).
450
Additional collections:
Norway: Nordland, Rana, Store Alteren, 5 collections; -
@rtfjellmoen, 6 collections; - Hammerness, map sheet 1927
I, VP 56, 5 collections; - Ramnaga, map sheet 1927 1, VP
56, 1 collection, T. Schumacher; - Rausandaksla, map sheet
1927 Ty VP 56, 1 collection; - Gronlbia; map sheet.20Z7aiy,
VP66,.. 2 1COLLECELOnN.
Sweden: Upland, Arentuna sn, Storvreta forest, N of the
railway station, 10 August 1942, S. Lundell (UPS); - V4ast-
manland, Sala, Grdéna gangen, close to Mans-Ols, 5 July
1945, R. Morander 89 (UPS).
Finland: Kuusamo, Liikasenvaara, lower course and mouth of
the brook Sirkkapuro, 24 August 1978 (4 collections, incl.
Fo. 7oeLO) and F.78.tl (C)); = duuma, “The Vuonas, 25 nuguee
LOLST (GC yar.
Estonian SSR: Rakvere, Neeruti, 4 September 1965, Raitviir
(TAAR AAT tC.
Switzerland: Graubtinden, E of the bridge over the river
Inn, ‘2 km SE ‘of, Ramosch, alt. L100 m,"6 ‘September y1o72,
Dissing: (GC).
France: Hautes Pyrénées, Le Chiroulet, Vallée de Lesponne,
sous’ Abtes pectinata, alt. 1100 m,” 21 duly 91974, Fo Can
doussau (C).
Italy: -Trento, Tertiolasi; in silva abiegna, Junto wloao,
leg. Bresadola, s.n. Humarta rhodoleuca Bres. n. sp. The
handwriting on the label is Bresadola's (Type, ex herb.
Bresadola, S$); - Sutdtyrol, in pinetis, 1884, leg. Bresa-
dola, s.n. Peztza permuda Cooke = Pez. patavina Sacc. The
handwriting on this label is not Bresadola's (ex herb.
Rehm, S).
U.S.A.: New York, Sagesville (= Lake Pleasent), August, C.
H. Peck (Type, NYS, NY, slide); - Lloyd-Cornell Preserve,
Slatersville, on bark of Betula lutea and very decaying
wood, 4 October 1947, C.T. Rogersen & R.P. Korft (CUP°3714g,
C); - Wisconsin, Perry's woods, Algoma, Kewaunee County and
vicinity, 16 August 1909, B.O. Dodge, s.n. Humarta rhodo-
leuca Bres. (NYS); - Alaska, Liberty Creek Trail near Chi-
tine, in wet leaf mold under alder, 1 September 1963, P.E.
Kempton # 9 (NY).
Figure 5. Rhodoscypha ovtlla.
SEM photo of spores. MO 72.22 (TRH). a. x 4000.
wee L200:
India: Chankaphug, Thimhu, Bhutan, on soil in mixed forest
of Blue Pine, Fir, Spruce and Rhododendron, alt. 9000 feet,
23 September 1980, R. Kaushal (PAN 18330, C); - Chuzom, Ha
Dzong, Bhutan, on soil infested with mosses and mixed with
Trorese “atter in. -aimixed. forest, alt. 10.000 feet, L Octo=
ber 1980,;'R. Kaushal (PAN 18364, C).
DISCUSSION
In the present investigation Humarta rhodoleuca Bres. is
considered to be identical with Peztza ovitlla Peck, with
the latter name having priority (cf. Dissing 1975).
Harmaja (1977), on the basis of spore characters, viz.
length, breadth, shape, content and thickness of the
episporium, distinguished between the two taxa. He also
paid attention to the substrate, H. rhodoleuca growing on
soll, and £. ovrlia, at least the material studied by hin,
growing on woody substrates.
However, the spore characters may vary considerably, even
in the same fruit body, and at present we are not able to
distinguish between two or more taxa. The results of some
spore-measurements are summarized in Table 1. The varia-
tion in the number of guttules is described above.
452
One collection from @rtfjellmoen (9 September 1975, (TRH) )
was pure white, but otherwise typical. After treatment by
an iron needle the touched and surrounding area changed
very quickly to rust red. The same feature could be demon-
strated on white specimens from Finland in 1978 (F.78.11
(C)).
Table 1. Spore-measurements in Rhodoseypha ovtlla.
10 spores measured in each collection (indicated in um).
Peztz2a ovtila, type, NYS 49.5-62.7 X 11.6-14.5
P. ovtlla, Alaska, Kempton # 9 36.3-45.5 X 10.6-13.9
Rhodoseypha ovtlla, Norway MO 75.61 33.0-46.2 x 10.0-16.5
Re OvULlag. Finland Fe) 78>10 34.7-43.0 X 11.6-14.0
Humarta rhodoleueca Bres., type, S 48.2-56.1 X 10.6-13.9
The specimens examined by Harmaja (l.c.) from Finland were
growing "on dead mossy logs of spruce (Ptcea abtes)". How-
ever, the 5 collections found by Dissing (see above), also
from the Kuusamo area in Finland, were fruiting on soil or
soil mixed with plant debris, like all collections from
Norway.
ECOLOGY
In Store Alteren, Norway (with 8 collections) Rhodoscypha
ovitlla is growing on a steep hill side in a mixed stanqpon
Picea abtes and Sorbus aucuparta with a very lush field
layer consisting of e.g. Aconttum septentrtonale, Lactuca
muralts, Ortganum vulgare, Victa sylvattca and tall ferns
(mostly Matteuceta). Hammernes (with 5 collections) is si-
milar whereas most collections in @rtfjellmoen (7 collec-
tions) have been made along the bed of an intermittent,
steep brooklet. The bed rock is limestone, the surrounding
vegetation is old, well-grown Picea forest with some
Betula.
In order to illustrate some edaphic parametres three soil
samples were taken. Collecting of the samples as well as
measurements of pH and conductivity in the samples are in
accordance with the methods described by Petersen (1967:
313-314). The results are summarized in Table 2.
453
Table 2. Edaphic factors for Rhodoscypha ovtlla.
Organic matter was determined as loss on igni-
tion after 6 H at 400 C. Methods for measure-
ments of pH and conductivity see Petersen (l.c.).
Collection pH LECUS) Loss on ignition
number (3 of dry weight)
MO 75.68 A 6.8 380 DGn/,
MON 5.6093 Gio 157 14.6
MO 73.73 ByAP 470 Siok
Rhodotarzetta Dissing & Sivertsen gen. nov.
Carposoma primum hemtsphaertcum, detnde cupulatum vel dt-
setforme, margtne prominente, base lata sesstle, extra
glabrum, roseolum ut semper hymentum, vel albtdum.
Everpuium —ommino textura intricata, cellults aehrots,
tineturam Cotton Blue tntra paulum acctptenttbus. Subhyme-
ntum e cellults dense tntertextts formatum, cellults unect-
ftets tineturam Cotton Blue avtde captanttbus tntermixtts.
Aset operculatt, non amylotdes. Paraphyses saeptatae, cel-
lula cugusque aptcalt guttulas extguas numerosas conttnente.
Sporae untsertatae, laeves, elltpsotdes, quaeque duas gut-
tulas continens.
Habttat in locts deustts.
Spectes typtfteca Rhodotarzetta rosea (Rea) Dissing & Si-
vertsen comb. nov., bastonyma Pustularta rosea Rea 1924,
Dowel.
Fruit bodies at first hemisphaeric, then cup-shaped to
disc-shaped, with a prominent margin; hymenium pinkish,
outside glabrous, concolorous with hymenium or whitish;
sessile on a broad base.
Pxcrpulum «throughout of. .textura intricata, ‘cells -colour=
less, with the content staining weakly blue in Cotton Blue.
454
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ie
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Figure 6. Rhodotarzetta rosea.
Marginal section.
x 130,
MO 75.81 (TRH,C).
456
Subhymenium of densely interwoven cells mixed with crozier-
forming cells, deeply staining blue in Cotton Blue.
Asci operculate, non amyloid. Paraphyses septate, uppermost
cell with numerous guttules.
Spores uniseriate, smooth, ellipsoid, with two guttules.
On burnt areas.
Type species Pustularta rosea Rea.
DISCUSSION
Rhodotarzetta rosea differs from species in the genus TYar-
zetta (Cooke) Lamb. in having an excipulum which throughout
is made up of textura intricata, in being glabrous, and in
possessing pinkish pigments in the paraphyses.
In Tarzetta species the excipulum is clearly two-layered
(cf. Eckblad 1968, Fig. 38 and Pant & Tewari 1970, Figs 2
and 17), the outside is furfuraceous, and all Tarzetia
species so far described have whitish, cream or greyish
pigments in the paraphyses. No Tarzetta species are con-
fined Lo Dune areas.
Rhodotarzetta rosea (Rea) comb. nov.
Pustularta rosea Rea, Trans. Worcestershire Nat. Club 8:20,
1924.
Tarzetta rosea (Rea) Dennis, British Ascomycetes 30, 1978.
Figure 6.
Fruit bodies solitary or in small clusters, 0.4-1.2 cm
broad, at first hemisphaeric, with a small circular opens
ing, then cup-shaped to disc-shaped, with a prominent mar-
gin; hymenium pinkish, sometimes with a purplish tinge,
when mature becoming pale orange; outside glabrous, con-
colorous with hymenium or whitish; sessile on a broad base.
Outer excipulum of textura intricata, 95-115 um thick, ina
dividual cells 3-8.5 wm broad, densely interwoven, sub-
parallel to outer surface, colourless, content staining —
blue in Cotton Blue.
457
Medullary excipulum also Of textura intricata, of loosely
interwoven cells, 5-11 um broad, content staining weakly
blue in Cotton Blue.
Subhymenium 30-65 um thick, of short, densely interwoven
cells, mixed with crozierforming cells, in which the con-
tent is staining deeply blue in Cotton Blue.
Hymenium 190-220 Um high; asci 11.6-13.2 um broad, opercu-
late, non amyloid.
Paraphyses septate, slightly enlarged above to 5 um broad,
uppermost cell with many small, reddish guttules.
spores uniseriate, smooth, ellipsoid, with two guttules,
ies ys Ito. Xx /.O=—6. 1-9 ..9 LM.
MATERIAL
Norway: Nordland, Rana, @Ortfjellmoen, map sheet 2027 IV, VP
86, burnt area in gravel pit, together with Octospora
rubens (Boud.) Moser, 9 September 1975, MO 75.81 (TRH,C).
The description is based mainly on the above collection.
In addition the following collections have been studied:
Denmark: Sjelland, Tureby, burnt area, 21 September 1964,
P.M. Petersen & H. Dissing (C); - Jylland, Klosterhede
Plantage, 8 km N of Struer, burnt area, 10 October 1969,
P.M. Petersen (C).
Holand: ~Noord-Brabant, Groet Peel, 27 April 1961, P.B.
Mansen, on burnt peat (L); - Drente, Dalen, 23 May 1961,
on burnt peat, J.J. Barkman (L).
458
Neotttella aphanodictyon (Kobayasi) Dissing, Korf & Sivert-
sen comb. nov.
Aleurta aphanodtctyon Kobayasi, Ann. Rep. Inst. Fermenta-
tion Osakar3s 9 7e1I67.
Leucoseypha borealts Eckblad, Nytt Mag. Bot. 15:52-53,
L960.
MATERIAL
Norway: Finnmark, Tana, Rustefjelbma, 22 July 1961, F.-E.
Eckblad, 61.93 (Type of Leucoscypha boreales, O); - Nord-
land, Rana, Virvassdalen, map sheet 2027 I, VP O05, naked
soil among scattered mosses, 12 August 1979, MO 79.15 (C),
79.96 (TRH); — tbids, 28 August 1981, 61.94) (TRE). = Joma
bru, map, sheet.2027'1I, VP 85, 31 July 1974 (TRH) ;9— Gouna.
vatn, map. sheet 202/111, VP +73, -in,dnundation (zone gc
reservoir 1) August) 19795 79.99.(7RH)e
Greenland: Pamiut, Frederikshab, 62° N72 49°06'W, among
Polytrtchum, 7 September 1979, T. Borgen 79.129 (C).
USS2A.: “Alaska, Barrow): “plot 419, US .IBP Tundvampeenme
Sites, FASTIN 156°41'E, on soil among mosses", 19 August
7¥974;'0.K. Miller & G.A. Laursen 11631\ ¢.n.Aleuri~asaiee
nodtetyon Kobayasi (CUP 58835); - Alaska, Barrow, within
walking distance from the Naval Arctic Research Laboratory,
on soil among Polytrtchum, 17 August 1980, H. Knudsen
(Kh) 22,C); = Hbids, Mo Lange: 4A 25,C);> = ibid. Lo suere
1980, RiP. Korf. (AK 30, CUP; A’ 297C); — 2D10.) 92 eae
1980, Dicsing ~& .Sivertsen § (A 41), 9A» 42,6; =) o1btoe,
B 6,9,10, TRH).
DISCUSSION
The collections from 1980 were all made during the First
International Symposium on Arctic Mycology held at Barrow,
Alaska from 16-23 August where also Dr. Kobayasi partici-
pated. Although it seems as if the holotype of Aleurta
aphanodtetyon is lost we feel sure about the identity of
this species. A. aphanodtetyon possess all characters typi-
cal for Weotttella species, viz. soil inhabiting, asso-
ciated to Polytrtchum species, paraphyses with carotinoid
pigments, guttulate spores with cyanophilic ornamentation,
spores and other cells with carminophilic nuclei, excipulum
of textura intricata, and with whitish flexuous or stiff
hairs on the outside.
A459
ACKNOWLEDGEMENTS
In 1972 the fieldwork expenses were largely covered by the
Royal Norwegian Society of Sciences and Letters. Grants
from the Norwegian Research Council for Science and Humani-
ties made later investigations possible. Part of the travel
expenses for H.D. was granted by the University of Copen-
hagen. 0. Mattsson, Institut for Planteanatomi og Cytologi
performed the TLC analyses, P. Milan Petersen, Institut for
@kologisk Botanik analysed the soil samples, J. Fuglsang
Nieleen, institut for Geologi og Palaeontologi, operated
the Scanning Electron Microscope. 7. Christensen prepared
Piommeiatin. diagnoses, i: Olson; Genetisk Institut A’ and
F.-E. Eckblad, Oslo gave valuable suggestions during prepa-
ration of the paper. K. Vestberg and W.H. Larsen prepared
the photographs. A.G. Kretborg typed the manuscript.
We greatly appreciate their co-operation.
460
LITERATURE
Arpin, N. 1969. Les caroténoides des Discomycetes: essai
chimiotaxinomique. -— Bull. Mens. Soc. Linn. de Lyon 38,
supplement 1-169.
Dissing, H. 1975. Peziza ovilla Peck = Humaria wrhodoltenea
Bres. >= ‘Bula; {Soceelvc wi Fr. Oleh ad.
Dissing, H. & S. Sivertsen, 1975. Operculate Discomycetes
from Rana (Norway) 1. Chalazton scetdbite, gen.aunovwe
Sp.y nov. aseNorwe dic, SOUS) (22:5 1—4%
Dissing, H. & S. Sivertsen, 1980. Operculate Discomycetes
from Rana (Norway) 3. Helvella rtvularts sp. nov. -
BOCs Ldsskee/ S25 LOL-102.
Eckblad, F.-E. 1968. The genera of the Operculate Disco-
mycetes. A re-evaluation of their taxonomy, phylogeny
and nomenclature. = Nytt. Mag. Bot.) dS: b-192%
Harmaja, H. 1977. Leucoscypha ovtlla n. comb., a species
new to Europe, found in northern Finland. - Karstenia
17: 73-76.
Pant, D.C. & V.P. Tewari, 1970. Observations on two species
of the genus Pustultna. - Mycologia LXII: 1187-1194.
Petersen, P.M. 1967. Studies on the ecology of some species
OfePezizaless = Bot. Tidsskr. .62.yolZ—s22e
Rifai, M.A. 1968. The Australasian Pezizales in the Herba-
rium of the Royal Botanic Gardens Kew. - Verh. Kon.
Ned: Akad. Wetensch., Afd. Natuurk.;, LE 573) 2 lo7
Svrcek, M. 1974. New or ‘less. known Discomycetes]) 2..—
eska Mykologie 28: 129-137.
MYCOTAXON
VOlexVie’ No. 2, pp. 461-499 January-March 1983
BASIDIOMYCETES APHYLLOPHORALES EPITHELOIDES ETALES
par J. BOIDIN et P.: LANQUETIN
Laboratoire de Mycologie associé. au C.N.R.S.
Université Claude Bernard, 43 boulevard du 11 novembre 1918
F.69622 - Villeurbanne - Cedex
France
The presence of sterile pegs is not a suitable character
to circumscribe a natural group. The fourteen species of warm coun-
tries studied (ten of which are new) are distributed among the genera
Epithele (Pat.) Pat. emend., Pteridomyces Jtilich emend. and Vesiculo-
myces Hagstrom emend. The main results are translated into English
at the end of the paper.
Patouillard (1899) propose une section Epithele dans le
genre Hypochnus, section caractérisée par des "émergences qui traver-
sent l'hyménium et qui sont comparables a celles de Mycobonia et de
Veluticeps dans le groupe des Stereum ou d'Heterochaete dans les
Hétérobasidiés". Cette proposition fait suite 4 la description
d'Hypochnus dussii qui est comparé a H. typhae.
En 1900, Patouillard éléve —Epithele au rang de genre, sans
désigner le type, comme cela se faisait 4 l'époque. Il cite les deux
mémes espéces et figure E. typhae. H. dussii ayant été transféré par
von Hohnel et Litschauer (1907) dans le genre Peniophora, E. typhae
est devenu, en fait, le seul représentant et a été logiquement sélec-
tionné comme type par divers auteurs (cf. Donk 1957).
Peu a peu des Epithele nouveaux ont été décrits par Bresado-
la, Burt, Rick, Cunningham et plus récemment par Boquiren (1971) dans
une révision ot l'on retrouvera citées toutes ces espéces. Mais comme
pour E. dussi qui est actuellement un Tubulicium, les émergences de
plusieurs espéces deprétendus Epithele sont en fait des cystides bien
différenciées enrobées d'hyphes gréles; ces espéces n'ont pas leur
place ici: E. capitata Rogers et Boquiren, E. macrospora (Liberta)
Boquiren, E. vermifera (Bourd.) Boquiren.
Ces exclusions faites, la révision de Boquiren comprend 8
espéces auxquelles il faudrait ajouter 4 espéces créées par Rick et
qui n'ont pas pu étre réétudiées depuis lors.
462
On peut s'attendre 4 ce qu'un regroupement basé sur un seul
caractére physionomique, la présence d'émergences stériles, crée un
ensemble artificiel; Patouillard avait d'ailleurs souligné lui-méme
des @émergences de méme erie chez les Hétérobasidiés et dans le
Pekoupe des Stereum"ce qui s'explique par un phénoméne de convergence
et n'est donc pas un témoignage de réelle parenté.
Nous traiterons ci-aprés de 14 espéces, récoltées en zones
intertropicales, 9 africaines, 2 malaises et 2 américaines, auxquelles
11 faut ajouter une espéce malgache trés particuliére avec ses spores
amyloides et ses énormes Gloeocystides sulfoaldéhydes positives que
nous ne pouvons éloigner du genre Gloeocystidiellum Donk sensu lato
malgré ses émergences stériles. C'est un exemple complémentaire de.
convergence!
Julich a successivement proposé 3 genres monospécifiques:
Epithelopsis (1976), Pteridomyces et Skeletohydnum (1979), sans parler
de Mycothele (1976) créé pour une espéce discoide que Patouillard
aurait assigné au "groupe des Stereum", et dont nous ne parlerons
pas Wel,
On peut dresser une clé schématique des genres étalés~-adhé-
rents en sélectionnant les caractéres distinctifs puisés dans les
descriptions génériques et spécifiques de Julich. Tous les représen-
tants sont pourvus de boucles et d'émergences stériles et, semble-t-i}l
d'éléments dendrophysoides.
— ‘Structure monomucique’ |:
. basides de 10-15 um de longueur; émergences faites
d'hyphes a paroi mince. Pteridomyces Jul.
type: E. galziniy Bres:
. grosses podobasides longues de 55-/0 um;
spores a paroi épaissie 4 maturité, non cyanophile.
Epithele Pat.s.str.
type select. : Es. typhae (Pers .jorac.
- Contexte dimitique, et @émergences faites
d'hyphes squelettiques.
spores a paroi épaissie et cyanophile, Epithelopsis Jul.
gloeocystides. type: E. fulva Cunn.
. spores a parol mince non cyanophile. Skeletohydnum Jul
type: E. nikau Cunn.
Aucune des 13 espéces a classer n'a de spores a4 parois cyano-
phile; elles devraient donc se répartir dans les genres Pteridomyces,
Epithele et Skeletohydnum en fonction d'une part de la taille des
basides, d'autre part du mitisme. Ce sont 1a deux critéres quantitatifs
ou progressifs. Le "mitisme' a été l'objet de bien des interprétations _
divergentes; c'est ainsi qu'Epithelopsis fulva (Cunn.) Julich est ~
considéré par le premier auteur comme monomitique, par le second comme
dimitique. Il faut rappeler que la notion de systémes d'hyphes mono-,
di-, et tri-mitiques est due a Corner qui décrivait ainsi dés 1932
la structure du contexte et des dissépiments de quelques polypores.
(voir la mise au point de Corner 1953 par exemple).
463
Corner ne prend en compte ni le cortex et les hyphes de la
face stérile, ni l'hyménium. Faut-il ou non prendre en considération
les 6mergences stériles des Epithele s. lato ? Il faut en outre bien
faire remarquer que la différenciation d'un deuxiéme systéme, fait
d'hyphes squelettiques, c'est-a-dire d'articles terminaux longs et
a paroi épaissie, peut @tre trés affirmée ou au contraire encore peu
marquée : ces deux cas ont été rencontrés dans notre étude. En fait nous
avons observé les situations suivantes :
a- contexte et émergences faits uniquement d'hyphes géné-
ratrices: cas d‘E. typhae et de P. galzinii, mais aussi
d'E. macarangae n. sp. one efibulata n.sp., et de P. lac-
teus n.sp. et sphaericosporus n.sp.
b- @mergences qui "naissant des parties profondes" comme
l'écrit Patouillard (1900 p. 59) sont faites en grande
partie d'hyphes squelettiques plus ou moins affirmées.
Entre les émergences, parfois trés rapprochées, le con-
texte ne montre que des hyphes génératrices (contexte
monomitique): cas du Skeletohydnum nikau et des E. gua-
delupensis, malaiensis et ovalispora ainsi que de P.
roseolus et P. sp. 8270.
c- é6mergences faites d'hyphes squelettiques,fibres nombreuses
dans le contexte entre les émergences (contexte dimiti-
que): ceci correspond 4 la définition du genre Skeleto-
hydnum mais non 4 son type; c'est par contre le cas
dq Bele. trispora:.
d- contexte dimitique 4 hyphes squelettiquesdominantes mais
émergences faites presqu'’uniquement d'hyphes de type
générateur associées 4 quelques hyphes axiales 4 tendance
squelettique : cas d'E. sp. 5697.
e- contexte dimitique et émergences faites d'hyphes de type
générateur: cas d'E. interrupta Bres.
Aprés ce constat on ne peut qu'hésiter a utiliser le miti-
sme pour des découpages qui seraient trés certainement arbitraires. Par
contre, on ne peut hésiter 4 répartir nos récoltes en deux groupes se-
lon la forme et la taille des basides. Les unes ont de petites basides
étroitement claviformes 14-20 x 3,8-5 um, atteignant 30 um de long chez
le seul P. lacteus, sans contenu remarquable (groupe I). Les autres
ont de grosses basides larges de 6-8 4 10-14 um plus ou moins brusque-
ment rétrécies a leur base, souvent plus larges dans leur partie moyen-
ne ou méme dans leur moitié inférieure, a (2)-4 forts stérigmates ar-
qués, et au contenu riche en lipides comme celui des spores (groupe II).
Remarquons encore une fois que dans le groupe I,qui devrait
correspondre au genre Pteridomyces, les hyphes sont soit du type "a"
(cas de P. galzinii, espéce type mais aussi de P. lacteus sp. nov.),
soit de type "b" (P. roseolus sp.nov. et surtout P. sp.n” 8270) ce qui
ne cadre plus avec la définition du genre Pteridomyces.
Tous les cas de mitisme se rencontrent dans le groupe II
meapl <\)1)-.
464
Contexte (racines des émergences exclues)
monomitique dimitique
Emergences q q
A/ sans hyphes typhae
E. effibulata
macarangae
E. interrupta
squelettiques
sp. 5697
S., nikau
B/ faites surtout | E. guadelupensis
E. malaiensis EB. sererispoce
d"hyphes squelet-| E. ovalispora
tiques
Tableau I. Champignons épitheloides du groupe II (a grandes
basides).
On pourrait envisager au moins deux découpages 1) en fonc-
tion du contexte (colonnes verticales), 2) en fonction de
la présence ou non d'hyphes squelettiques dans les émer-
gences (ligne A: Epithele, ligne B: Skeletohydnum). Sont
soulignées les espéces types.
Ces constatations peuvent amener deux types de décisions
opposées:
1°- se baser sur la diversité des cas rencontrés pour poursuivre 1'écla-
tement et, par exemple, créer un genre pour les Pteridomyces du type
wo
"bp", les Skeletohydnum du type "c", ceux du type "e"... Si l'on veut
en outre tenir compte des particularités des parois sporiques, des
boucles... on peut envisager un plus grand nombre encore de combinai-
sons donc de genres nouveaux trés homogénes mais souvent monospéci-
fiques (mais alors 4 quoi servent les genres?),
2°- se baser sur cette méme diversité pour refuser la multiplication des
genres et accepter que les tendances évolutives, toujours en cours,
provoquent une certaine diversification, donc une certaine hétérogéné-
ité a l'intérieur des genres concus plus vastes mais constitués d'espé-
ces que l'on suppose naturellement apparentées.
Nous choisirons la prudence, c'est-a-dire la deuxiéme solu~
tion et ne créerons aucun genre. Le genre Pteridomyces sera ouvert aux
espéces a4 émergences de nature squelettique.
Pour le groupe II, trois alternatives se présentent: définir
4 genres, n'en retenir que deux qui correspondraient soit aux deux
colonnes verticales (l'un 4 contexte monomitique, l'autre 4 contexte
dimitique) soit aux deux lignes horizontales (émergences faites ou non
d'hyphes squelettiques:Epithele et Skeletohydnum ; la troisiéme solu-
tion consiste comme pour le groupe 4 petites basides a ne retenir qu'un
seul genre et donc 4 réduire Skelethohydnum en synonymie d'Epithele.
465
Ceci revient 4 reconnaitre deux lignées, l'une aux petites
basides claviformes et aux spores de moins de 9 um de longueur, l'autre
aux grosses basides et aux spores de plus de 10 um de longeur, chaque
lignée ayant plus ou moins évolué vers le dimitisme. Seule
l'étude d'un plus grand nombre d'espéces, récoltées en des points les
plus divers du globe terrestre permettra de mieux préciser les ensem-
bles génériques naturels 4 distinguer.
En attendant, nous décrirons ci-aprés, dans l'ordre, les
espéces
1. du genre Epithele, 2. celles du genre Pteridomyces emend.,
et enfin 3. le Vesiculomyces épitheloide.
Pour chaque espéce, dans la mesure du possible nous ajoute-
rons les caractéres culturaux qui seront en outre résumésselon le
code de Nobles déja complété par J. Boidin (1966 a p. 6) et P. Lanquetin
Mo7sapsio7 note. 2);
D'autres additions paraissant utiles, nous regroupons ci-
aprés,avec ces nouvelles propositions l'ensemble des compléments antérieu-
rement publiés.
Les numéros soulignés sont ceux qui sont proposés pour la
premiére fois.
Oxydases
2: laccase et tyrosinase présentes simultanément
2a: laccase seule présente
2b: tyrosinase seule détectée
Boucles
~
3c: boucles constantes (présentes 4 toutes les cloisons
vraies)
31: boucles inconstantes (des cloisons simples séparent
certains articles nucléés)
3r: boucles rares
Eléments dextrinoides
8d: fibres dextrinoides (ex: Scytinostroma)
25d: dichophyses dextrinoides (ex: Vararia, Dichostereum)
28: asterophyses dextrinoides (ex: Asterostroma), P. Lanquetin
avait en 1973 proposé pour celles—ci le n°19, préoccupé
par Nobles (1967) pour "des cellules terminales
moniliformes"
Gloeocystides
15: signifie présence de gloeocystides sans précision sur
la réaction du conténu. Nous proposons d'ajouter
15a : gloeocystides sulfo-aldéhydes positives
15b : gloeocystides sulfo-aldéhydes négatives (Boidin et al.1980)
: gloeocystides avec schizopapille(s)
Thallie
59: signifie bipolaire (monofactorielle), 60:tétrapolaire
(bifactorielle),
59A: amphithalle bipolaire
60A: amphithalle tétrapolaire (proposé in Boidin et al.1976
Da 6258)
Comportements nucléaires au cours du cycle
61: normal: spore uninucléée, mycélium primaire (=myc. I),
uninucléé, mycélium secondaire (myc. II) binucléé.
62: subnormal: spore binucléée, myc. I agé uninucléé, myc. II
binucléé.
466
~
63: hétérocytique : myc. I 4 articles terminaux plurinucléés
myc. II binucléés
64: myc. I inconnu, myc. II binucléé
65: astatocénocytique: myc. I multinucléé, myc. II binucléé
ou multinucléé selon les conditions d'aération; marge
formée d'hyphes axiales multinucléées. (va généralement
de pair avec le caractére 4)
66: holocénocytique: myc. I et II multinucléés,
67: holomonocaryotique (Ktihner 1977): mycélium (monosperme
comme polysperme) uninucléé
68: holodicaryotique (Kthner 1977): spore binucléée germant
en un mycélium binucléé.
Clé des champignons épitheloides étalés des régions intertropicales
A- Spores amyloides réniformes, 7-8,5 x 4,5-5,5 um; grosses gloeocysti-
des 150-250 x 10-20 um sulfoaldéhyde-positives. Basidiome saumoné
a bordure blanche. Madagascar.
14 Vesiculomyces epitheloides
A- Spores non Sh EE stellar is sale ays Salita eee ase Sales eae elon seis hemi rece nk
B-Spores petites n svaveueiant pas 9 x 5 um; petites basides claviformes
15-20-(30) x 45m: sPteridomyces -emend s../2 05a. Aas oe: Severe tan ELA
B- Spores et basides plus grandes: Epithele emend: i. ..2.2.53. ose mens F
C- Spores subsphériques 4,5-5,5 x 3,8-4,8 um, basidiome blanc gris&atre,
émergences faites d'hyphes bouclées a paroi épaissie. Céte d'Ivoire.
13 P. sphaericosporus
C- Spores oblongues, ellipsoides ou subcylindriques.......... oa haere
D- Basidiome blanc de craie, émergences faites d'hyphes a paroi mince.
Spores ellipsoides 6-8 x 3-4 um. Gabon.
10 P. lacteus
D- Basidiome beige ou beige rosé, é6émergences faites d'hyphes 4 paroi
epaisse rét ievyanophile. is. sees Sia elahetoons oS Sieiccsle, aw -eventatetmenns E
E- Spores stubcylindriques5-6 x 2-255 ‘um;
République Centrafricaine. 11 P. roseolus
E- Spores oblongues 3,5-4 x 2;5-2,8 jm
Singapour. 12 P. sp. 8270
F- Emergences faites uniquement d'hyphes 4 paroi mince...... ole ite e eoae are Ge
F- Emergences faites d'hyphes de type squelettique, raides, serrées
en faisceau, parfois recouvertes d'un manchon d'hyphes bouclées...I
G- Pas de boucles; spores fusiformes, 17-21 x 7-10 um, sur Palmae;
Gabon, Sierra Leone (7?) 2 E. ef£ibulata
G- Boucles; spores ellipsoides, 14-20 x 6,5-8,2 um, émergences obtuses
fragiles; République Centrafricaine..... aie esate star SUR a sha et aeeerae sien
H- Contexte monomitique sans fibres. 5 E. macarangae
H- Contexte nettement dimitique, Zaire. 4 E. interrupta Bres.
I- Spores nettement biapiculées, 14-18 x 7-8 um; contexte dimitique;
Céte d'Ivoire, Gabon. Lie Crerispora
I- Spores ellipsoides, oblongues ou ovoides....... see Aree wate creeoeet re aC:
J-uSporesilarges derplus; devs: wate... Was es. sa ehete Jule Peisid ous eee rg
J- Spores ellipsoides ne dépassant pas 6 wm de largeur.........-.+-..L
K- Spores oblongues subelliptiques 17-21 x 8-10 um; sur Palmae;
Singapour. 6 E. malaiensis
467
K= Spores ovoides, 13,5-17,5 x 8,5-11 um; sur Palmae;
Gabon. 8 E. ovalispora
L- Contexte pais, lache, riche en fibres; champignon détachable. Spores
ellipsoides 10-12 x 5-6,5um; Argentine.9 E. sp. 5697
L- Contexte mince et dense; champignon trés adhérent...... Bias ce alin
M- Spores 10-14 x 4,5-6 um; Gabon. 7 E. nikau Cunn.
M- Spores 14-18 x 4,5-5,8 um; Guadeloupe 3 E. guadelupensis
I - Genre Epithele (Pat.) Pat. 1900.
Synonyme: Skeletohydnum Julich 1979
Nous placgons ici les espéces 4 grosses basides et grandes spores
blanches ou crémes, 4 paroi mince ou un peu épaissie, lisses et qui ne
sontni amyloides, ni dextrinoides, ni cyanophiles. Les hyphes généra-
trices sont avec ou sans boucles. Les hyphes squelettiques sont absentes
ou présentes dans le contexte, dans le contexte et les émergences, ou
dans les @mergences seules.
l- Epithele citrispora Boidin, Lanquetin et Gilles, sp. nov.
Bais. yl
Jacens, solida, adhaerens, alba vel cremea, emergentiis concoloribus,
regulariter dispositis hirta, 40-50 um crassa. Trama hyphis skeleticis
permultis (praeter in parte infima). Emergentiis 75-170 x 30-80 um
transgredientibus, ex hyphis skeleticis verticalibus, tunica crassissi-
ma, praeter in superiore parte, quae speciem dendroideam accipere
possunt, constantibus. Basidiis 35-50 x 9-12 um, fibulatis. Sporis
biapiculatis, 14-18 x 7-8 um, haud amyloideis neque cyanophilis, guttu-
lis oleaginosis completis. In ligno. Holotypus LY 8537. Gabon.
A la récolte, largement étalé, solide, adhérent, blanc un peu grisa-
tre (Munsell 2,5 Y 8/1 a4 10 YR 8/1), tout couvert de petites aspérités
concolores; marge similaire assez brusque, parfois pruineuse.
En herbier, ¢talé gris clair quand il est trés mince (5 Y 8,5/2) a
ereme (5 Y°8,5/3,5) ou. creme sale (2,5 Y 8,5/4), adhérent 4 marge brus-
que ou au contraire progressivement amincie et plus blanche; sous la
loupe il est tout hérissé d'émergences concolores assez réguliérement
disposées.
Epais de 40-100-250 um, il est formé d'une couche inférieure dans
laquelle prennent naissance les émergences entre lesquelles s'établit
une couche supérieure hyméniale bien différente. La couche inférieure
haute de 15-160 Um peut montrer contre le support quelques hyphes géné-
ratrices bouclées a4 paroi un peu épaissie, collées entre elles en une
sous-couche horizontale dense de 5-15 um d'épaisseur; trés vite s'y
mélent des hyphes squelettiques larges de 2-2,8 um non cyanophiles, 4a
lumen subnul, les plus profondes horizontales, mais vite entrecroisées
et emmélées en tous sens, parfois ramifiées, qui forment l'essentiel
de la couche inférieure; les hyphes génératrices 4 paroi mince ou peu
épaissie, x 1,5-1,.8-(3) um semblent trés rares.
Emergences subcylindriques ou un peu cOniques mais obtuses, ancrées
a la base du basidiome et dépassant 1'hyménium de 75-170 x 30-50-80 um;
elles sont formées d'hyphes de type squelettique dressées parallélement
en faisceau; elles sont larges de 2,5-(3,8) um, leur paroi trés épaisse
s'amincit 4 l'extrémité qui peut se ramifier et prendre un aspect den-
droide; pas ou peu de cristaux dans les émergences.
Figure 1: Epithele citrispora sp.nov. : a- hyménium, et sl1,spores
(holotype LY 8537); b- contexte, et c- sommet d'une émer-
gence (LY 7158); s2- spores (LY 9013).
Les échelles sont valables pour les figures 1 a 8.
469
Hyménium formé de basides trapues, 35-50 x 9-10-12 um, brusquement
rétrécies A la base qui est bouclée, 4 4 forts stérigmates 7-14 x 3 um;
leur contenu est guttulé gras; elles sont portées par des hyphes sous-
hyméniales bouclées trés ramifiées qui peuvent émettre des hyphes den-
droides A paroi mince, souvent peu remarquables, qui atteignent la
surface; dans le sous-hyménium peuvent s'engager des hyphes squeletti-
ques verticales issues de la couche inférieure, qui montent plus ou
moins haut et peuvent émerger isolément ou en groupes.
Spores citriformes biapiculées, 14-18 x 7-8-(9) um, non amy loides
ni cyanophiles au contenu multiguttulé. Pour lestypesaxs= 15,17 27170
Ree O25 aour, 9013, % = 116,06. 1, 1" x 7,95 %-0;41.
C'est la seule récolte africaine qui réponde tout a fait 4 la dia-
gnose du genre Skeletohydnum Julich. Elle ne peut cependant pas étre
séparée génériquement d'E. ovalispora, fusispora, malaiensis...etc.
Récoltes: COTE D'IVOIRE: LY 7158, forét du Téké prés d'Abidjan, 29
AVTL) 1OVS-G.eGilles 210.
GABON: LY 7843, sur branchette, ile aux chauves-souris,
Makokou, 13 mai 1976 (*): LY 8537, HOLOTYPE, sur bois dans
le fourré littoral, N. de Libreville, 21 janvier 1978, G.
Gilles 1070; LY 9013, en forét ombrophile, ibid., 3 décem-
bre 19/78,.G. Gilles 13203.LY 9106, fourré 1ittoral,. 6 jan-
vier 1979; LY 9142, forét ombrophile, 27 janvier 1979 et
LY 9199,, ibid., 17 février 1979, tous trois leg. G. Gilles
sans n°.
Caractéres culturaux d'E. citrispora
SPORES.- (LY 7158-8537-9013-9106) uninucléées, mais pour un grand nombre
de spores, le contenu reste trés hétérogéne et il est impossible de
préciser.
GERMINATIONS.- Elles n'ont pas été obtenues malgré plusieurs essais
avec LY 7158,..7/843, 9013 et 9106.
POLYSPERME.- Seul LY 7158 s'est développé.
Croissance: trés rapide (boltes couvertes en deux semaines)
Aspect: La marge est réguliére. Le mycélium présente un aspect fidéle
dans toutes les boites: la surface de la culture est uniformément lisse,
blanc pur, mate 4 jour frisant. Ce disque blanc est souligné, 4 la
périphérie contre le verre, par une fine ligne presque continue brun
sombre (vers5 YR 3/3 chocolat)) et il peut aussi trés localement mon-
trer de minuscules points brunatres (5 YR 4/6 et 3/3). Dans l'une des
boites le mycélium teinté forme une petite plage (fawn R., 5 YR 5,5/4).
Aprés trois mois une autre bofte montrait plusieurs petites plages
brunes et des zones densément piquetées de petits points bruns.
Revers des cultures: inchangé. Odeur nulle.
Microscopie:
Mycélium aérien: il forme une peau détachable 4 la pince mais assez
cassante. Cette peau est constituée surtout d'hyphes réguliéres, x 1,2-
2 um, 4 paroi mince, bouclées, densément emmélées avec des hyphes plus
larges x3-4 um, du cdté de la gélose. Vers la bouture, quelques hyphes
superficielles portent des cristaux réfringents pouvant former, une gai-
ne. Dans les petits points bruns, les hyphes, irréguliéres, ont une
parol nettement brunie qui s'épaissit aux extrémités.
* s:quand le récolteur est J.Boidin,il n'est pas indiqué.
470
Mycélium submergé: hyphes a4 paroi mince, bouclées, x 2-8 um, riches
en gouttes réfringentes, les étroites bien cylindriques et les élargies
plut6t irréguliéres avec boucles surbaissées. Quelques hyphes régulié-
res, x (2,5)-4-7 wm, ont une paroi nettement épaissie, x0,5-1 um,
prenant mal le rouge Congo, elles sont bouclées et montrent aussi de
nombreuses cloisons de retrait bombées.
Boucles: constantes.
Cytologie: Hyphes aux articles réguliérement binucléés, bouclés.
Oxydases: ac. gallique: +++++,0 gaiacol: ++++,0
p.-crésol: -, précipité blanc tyrosine: -, 10 mm
CODES: 2acay oC 32 BG 36a = 425 = 54 = 64
Remarque. A part la formation d‘unfin liseret brun autour des cultures
trés blanches, et la forme spéciale des boucles sur les hyphes submer-
gées, le mycélium de cette espéce est trés peu caractéristique.
2. Epithele efibulata Boidin, Lanquetin et Gilles sp. nov.
Figw 2
Parvis maculis formata, deinde membranacea, alba vel cremea, emer-
gentiis gracilibus, concoloribus praedita.Trama monomitica, sine fibu-
lis. Emergentiis ex hyphis horizontalibus cohaerentibus, tunica tenui
vel parum incrassata constantibus. Basidiis 32-52 x 8-10 um. Sporis
ellipsoideis, 17-21 x 7-10 um, tunica paulum incrassata ac aetate luteo-
la, haud cyanophila neque amyloidea. In Palmaceis. Holotypus LY 7863.
Gabon.
A la récolte, il se présente en petite taches distinctes puis conflu-
entes; il est membraneux, détachable sous l'aiguille, blanc puis créme
(2,5 Y 8/2 a 8,5/4), créme alutacé (10 YR 8/3 et méme 8/4) avec marge
lisse, blanc grisatre (2,5 Y 8/2 ou 10 YR 8/1,5); sous la loupe il est
couvert d'émergences gréles, concolores.
En herbier, il apparait mince, adhérent, fragile (pulvérulent sous
le rasoir), parfois fendillé, alutacé pale (10 YR 9/3,5 9/4) 4 créme
(2,5 Y 8/4) avec marge amincie plus pale et sans émergences.
Epais de 100-200 um hors des émergences qui débordent de 70-180 x
25-35-(50) um. Couche basale, haute de 15-50 um, d'hyphes horizontales
serrées, x 2-3 wm, 4 paroi mince et cloisons simples, puis hyphes obli-
ques redressées formant une zone transitoire plus lache: ces hyphes,
x 2-2,5 um a paroi mince, sont fréquemment ramifiées et peuvent contenir
des gouttelettes lipidiques; dans cette zone se déposent parfois d'abon-
dants amas cristallins. .
Emergences constituées d'hyphes paralléles, cohérentes, a4 paroi mince
ou peu épaissie, sans boucles; a leur surface on peut voir des extrémi-
tés dendroides; dans la partie profonde de ces émergences on peut trou-
ver de rares hyphes 4 paroi plus épaissie et des dépéts engainants de
cristaux détersiles.
Zone hyméniale dense, formée d'hyphes gréles, x 2 um, et de basidio-
les claviformes riches en huile entre lesquelles peuvent émerger des
hyphes dendroides (visibles surtout dans le bleu lactique) dans lequel
on voit en outre des dépéts de fins cristaux bacillaires sur les élé-
ments superficiels. Basides 32-52 x 8-10-(11) wm au sommet mais plus
large 4 mi-hauteur (par ex. 13 um) portant 4 gros stérigmates longs de
9-13 wm; elles peuvent émerger de 12-20 um au-dessus des basidioles.
ATi
Spores fusiformes (14)-17-21-(23) x (6)-7-10-(11) wm, @ sommet par-
fois un peu pincé (tendance biapiculée), au contenu gras, 4 paroi fai-
blement épaissie (moins de 0,5 um). Cette paroi n'est ni amyloide , ni
cyanophile. Sur les exsiccata quelques spores semblent avoir une
paroi épaissie mais c'est une impression due 4 la rétraction et au dé-
collement du cytoplasme; ces spores A4gées peuvent apparaitre jaunatres
et flétries, elles ont une section polygonale.
Les mesures statistiques sur sporées vont de 17,23 x 7,26 pour 9146
a 20,57 x 9,28 pour 9155 mais i1 s*agit dans ce‘dernier cas d'im basi
diome largement étalé, alors que 9146, jeune encore, est formé de
petites taches disjointes.
Récoltes: GABON: LY 7863, HOLOTYPE, sur Raphia sp., marigot a Makokou,
17 mai 1976; LY 7867 ibid.; LY 9120, sur Eremospatha macro-
carpa, La Mondah, 2] janvier 1979, G.. Gilles {140/72 LY 9145
et 9146, sur Elaeis guineensis, La Mondah, 3 février 1979,
G. Gilles 1425 et 1426; LY 9155 sur Raphia sp., 10 f£évrier
1979, G. Gilles 1436; LY 9167 et 9168, sur Elaeis guineensis,
18 février 1979; G. Gilles 1447 et; 144683) LY 9182) ibide
24 février 1979, G. Gilles 1459; LY 9206 ibid... (10: mare
1979, G.9Giltes* 14793" LY 9277, abid.4) T4uavrialp loi. aGe
Gilles sans n
C'est la seule espéce d'Epithele qui soit dépourvue de boucles. Nous
avions pensé, a la lecture de Boquiren que cette espéce pourrait étre
Epithele interrupta Bres. Il faut abandonner ce point de vue aprés étude
du type (in BR) qui vient du Zaire (Mayidi, ler février 1910 leg.
Vanderyst). Il s'agit d'une récolte avec émergences dispersées, peu
nombreuses, courtes, obtuses, fragiles, faites d'hyphes bouclées 4
terminaisons dendroides, mais dimitique par des fibres hyalines nom-
breuses dans le contexte. Boquiren avait étudié une récolte de Sierra
Leone déterminée E. interrupta (in DAOM 99.845, sur Elaeis guineensis)
qui ne peut étre l'espéce de Bresadola; bien que Boquiren ne précise ni
la présence ou l'absence de boucles, ni celles d'hyphes squelettiques,
certaines de ses données et figures laissent 4 penser qu'il pourrait
Sagi nd hecveribulata.
Caractéres culturaux d'E. efibulata
SPORES.- (LY 7863, 7867) uninucléées.
GERMINATIONS.- non obtenues
POLYSPERME.- (LY 7863, 7867, 9120)
Croissance: trés rapide (boites couvertes en deux semaines)
Aspect: la marge est trés réguliére. Le mycélium aérien blanc pur forme
une peau feutrée, appliquée, lisse, mate, pouvant toutefois montrer
un ou deux plis. A la périphérie des cultures 9120, il apparait grume-
leux-floconneux et se teinte localement de créme 10 YR 9/2. A la péri-
phérie de 7867, il se forme de petites masses de 1 4 8 millimétres de
diamétre hérissées d'aiguillons un peu jaunatres, et exsudant des
gouttelettes hyalines a ambre clair.
N.B. Les cultures LY 9120 révélent par transparence des plages un peu
rosatres, pratiquement non discernables dans les boites observées par
dessus ou par dessous.
Le revers des cultures est inchangé, l'odeur nulle.
ATS
Microscopie:
Mycélium aérien: il est constitué d'hyphes réguliéres, x 1,5-2-(3)
um, a parol mince, sans boucles. Beaucoup d'hyphes sont affaissées. .La
peau feutrée prélevée dans la partie agée, est formée d'éléments soli-
dement agglomérés, trés difficiles 4 séparer par écrasement; il s'agit
d'hyphes, x 2-3-(5) wm, sinueuses, contournées, ramifiées, emmélées,
a paroi hyaline épaisse 4 trés épaisse, (1,2-1,5 um par ex.) formant
un puzzle dense. Les plages légérement colorées vues par transparence
sous le mycélium superficiel blanc de LY 9120, correspondent 4a une
crotite coriace pareillement constituée en puzzle. Les émergenges obser-
vées seulement dans LY 7867 sont formées d'un faisceau d'hyphes paral-
léles, x 1,5-2-(2,5) um, réguliéres 4 paroi mince, sans boucles. Toute-
fois quelques rares hyphes ont une paroi nettement épaisse. Certaines
hyphes 4 paroi mince, sans boucles, se ramifient vers leur extrémité
mais la présence de petits cristaux géne liobservation. 1] faut utili-—
ser le mélange KoH-Phloxine pour bien voir ces courtes ramifications
a aspect de fines dendrophyses.
Mycélium submergé: Aprés avoit 6té la peau blanche du mycélium
a@rien, on peut observer de rares lignes blanches sillonnant le mi-
lieu gélosé. Une telle ligne bien visible sur une coupe transversale
apparait faite d'hyphes x 2-3 um, a paroi épaisse trés intriquée en
puzzle.
Ailleurs le mycélium submergé est formé d'hyphes x 4-6 um, assez
reguiueres,. a) cloisons’ simples; et de rameaux, x 1,5=2 jm, assez
ramifiés. Quelques hyphes, x 3-4 um, ont une paroi épaissie a nettement
Epaisse, x 0,8-1 jim, ce ne sont. pas. des. fibres.
Boucles: absentes
Cytologie: les trois polyspermes sont formés d'hyphes aux articles
réguliérement uninucléés. Le croisement des deux polyspermes 7863 et
7867 a donné un mycélium aux articles également uninucléés. L'espéce
pourrait étre parthénogénétique.
Oxydases: ac. gallique +++++,0 garacoL: +++++-.0
peeeresole =o precapite, blanc tyrosine:) =, .0*4a<20. mn
CODES a ea Ome mS 2 <=. 360 3G. G2— 54 = 67
Remarque. Cette culture est faiblement caractérisée par sa croissance
rapide, la présence d'hyphes agglomérées en puzzle et par ses cultures
polyspermes sans boucles aux articles uninucléés. Cette espéce holomono-
caryotique est vraisemblablement parthénogénétique haploide ce qui
explique l'absence de boucles. De telles espéces ne peuvent é@tre séparées
généralement des espéces parentes binucléées. C'est le cas par exemple
d'Hypodontia efibulata Erikss. et Hjortst., (Hassan et David, inédit),
d'un Podoscypha centrafricain sans boucles (inédit) de Vararia auran-
tliaca Boid. et Lanq. (Boidin et al 1980).
3. Epithele guadelupensis Boidin et Lanquetin sp. nov.
Fig. 3"A
Jacens, tenuis, fragilis, ex alba griseola, im margine levi pallidior,
30-50 um crassa, emergentiis albis, regulariter dispersis, 75-130 x 15-
20 um, crassamentis crystallinis vestitis, ex hyphis rigidis, refringen-
tibus, tunica incrassata. Hyphis genetricibus fibulatis, stratum densum
pseudoparenchymaticum formantibus, deinde in perpendiculum directis.
Basidiis 22-35 x 6-7,5 um, 4 sterigmatis divergentibus; adsunt nonnullae
hyphae dendroideae inter basidiola et in emergentiis. Sporis ellipsoi-
deis, 14-18 x 4,5-5,8 um, haud amyloideis neque cyanophilis. In Palma-
ceis. Holotypus LY 8119. Guadeloupe.
474
A la récolte, étendu, mince, blanc grisatre (neutral 8/0) avec marge
atténuée pruineuse sous la loupe, lisse sur un demi m puis 4 petites
émergences blanches réguliérement disséminées.
En herbier, étalé, adhérent, blanc grisatre (neutral 8/0) puis créme,
atteignant (5 Y 8/3,5) avec marge amincie plus blanche; les émergences
apparaissent trés t6t prés de la bordure et s'allongent ensuite, elles
sont nombreuses, serrées. Sous le rasoir le champignon est fragile et
cassant.
En coupe, trés mince 30-50 um environ avec émergences élancées, cdéni-
ques, 75-100-130 x 15-20 pm, recouvertes de dép6ts cristallisés, prenant
naissance 4 la base du basidiome. Sur quelques hyphes génératrices bou-
clées A paroi mince, formant une mince couche d'aspect paranchymateux
en coupe transversale, se dressent les éléments du sous-hyménium et de
1"hyménium.
Emergences formées d'hyphes raides réfringentes x 1,8-2,2 um, 4a
paroi épaissie sauf vers l'extrémité effilée, assez congophile aprés
traitement ammoniacal 4 60°; elles portent souvent en surface des hyphes
a parol mince qui peuvent se redresser et montrer des terminaisons
dendroides; a leur base les hyphes 4 paroi épaissie sont sinueuses par-
fois ramifiées.
Basides trapues, souvent un peu étranglées, 22-35 x 6-7,5 4 4 stéri-
gmates de 7-8 x 2 um, @écartés. Des hyphes dendroides émergent entre les
basidioles au contenu guttulé. he
Spores ellipsoides de face, 14-18-(20) x 4,5-5,8 um, (x = 15,97 2
1,20 x 5,18 + 0,32), au contenu gras, 4 paroi mince non amyloide ni
cyanophile; l'apicule est réfringent.
Récoltes: GUADELOUPE: LY 8119, HOLOTYPE, sur rachis de Prestoa montana
(Palmae) au sol, Bras David, 4 octobre 1976.
Caractéres culturaux d'E. guadelupensis
SPORES.- (LY 8119) uninucléées.
GERMINATIONS.- elles sont prélevées deux jours aprés la dispersion
MONOSPERMES.- A trois semaines, sur 22 prélévements, 13 sont bouclés,
3 montrent localement des crochets et 3 sont a cloisons simples. La
coloration cytologique de ces derniers a donné pour deux d'entre eux
des plages totalement binuclééeset des plages aux articles réguliérement
uninucléés. Le troisiéme, moins développé, s'est révélé constitué d'‘ar-
ticles trés réguliérement uninucléés sans boucles, un mois plus tard,
cette méme culture était formée d'hyphes aux articles réguliérement
binucléés bouclés. L'espéce est donc homothalle.
POLYSPERME.-—
Croissance:trés rapide (boites couvertes en deux semaines)
Aspect: marge réguliére. Mycélium aérien blanc pur d'aspect typique,
faisant penser 4 une surface farineuse laquelle aurait été balayée pres-
que totalement a certains endroits. Il est plus dense dans la zone jeune
ou de nombreuses fibrilles entrecroisées forment un réticule peu mis en
évidence car rigoureusement blanc sur fond blanc. A la périphérie, les
fibrilles, trés fines, grimpent contre le verre et vont, en s'étalant
décorer la face inférieure des couvercles.
Dans une bofte, nous avons observé de trés petites plages crayeuses,
de 1,5-2 millimétres de diamétre, ce sont des fructifications qui donnent
une sporée créme sale dans le couvercle. Le revers des boites est inchan-
gé, l'odeur nulle.
475
v2,
fe
X,
SEG
Ss >
Sw
&f
ae SD,
belgie:
(holotype LY 8119).
is Sp.nov.
Brese:
uade Llupens
Big [NS Epithele
Breune
s sur le type,
spores vue
Epithele interrupta
et deux hyphes dendroides.
oe
476
Microscopie:
Mycélium aérien: il est canstitué par des hyphes réguliéres étroi-
tes, x 1,2-1,75 ym, “4 parol mince, \boucles constantes) (en fait
beaucoup sont des khyphes collapsées) et quelques hyphes plus
larges, x 4-5 um, réguliéres boucléées 4 paroi mince et contenu
homogéne.
Fructifications:elles montrent de nombreuses émergences pointues de
100 3 140 um de haut, de nombreuses basides 27-35 x 6,5-7 um au
sommet, boucléés 4 leur base étroite, x 2 um, et portant sur 4 stéri-
gmates longs jusqu'a 8 um, des spores allongées 15 x 5 um.
Mycélium submergé: Dans le milieu, juste vers la surface, hynhes
x 2,5-4 um, boucléés, 4 paroi mince, trés irréguliéres avec multi-
ples bosses ou courtes excroissances; des rameaux trés gréles, x l-
2 ym, sont aussi trés irréguliers. Quelques axes, x 3,5-5 um, sont
réguliers, a paroi @épaissie, congophile (¢€X- 0,5 sur hyphes de 4,1,
1 y sur hyphe de 5 y ), et portent des boucles en tibia.
Quelques rameaux bouclés trés tortueux comme s'ils allaient s'im-
briquer en puzzle mais sans y parvenir, ont une paroi ferme, trés
réfringente, qui ne prend pas le Rouge congo.
Boucles: constantes
Cytologie: hyphes aux articles binucléés, bouclés
Oxydases: ac. gallique: +++++, tr. gaiacol: ++++, 0
ps-verésol:e=, Léger précipité blanc. tyrosine: —7 tr.
CODE S024 = 360 a3 20= SO seoo) -42. alts oa 2 Oe od
Remarque. Culture peu caractérisée, sinon par son aspect (mais nous
n'avons qu'une souche!) et son comportement homothalle lent similaire
a celui d'E. nikau.
4 - Epithele interrupta Bres. ap. de Wildeman
Bull. Jard. Bot. Etat (Bruxelles) 4: 25, 1914
non E. interrupta sensu Boquiren 197]
Fig. 38
En herbier, étalé, mince, membranuleux fragile, créme, 4 marge
amincie blanchatre. Emergences peu nombreuses, courtes, irréguliérement
disséminées, détachables et laissant une cicatrice circulaire.
Epais de 90-150-250 um avec hyphes basales distinctes, assez hori-
zontales, réguliéres, x 2-3,5 um, bouclées, a paroi mince; s'y ajou-
tent bientdt de nombreuses fibres hyalines x 1,5-3 um 4 paroi trés
épaissie, s'arrétant a la base de l'hyménium. Le contexte peut accu-
muler des amas de cristaux. Hyménium fait de basidioles (environ 60
x 11 um) au contenu gras passant 4 des basides larges de 11-13 um
a 4 gros stérigmates; des hyphes dendrophysoides parfois nombreuses.
Les émergences, 200 x 150 um, sont entiérement constituées d'hy-
phes bouclées a paroi mince ou presque, x 2-4 um, parfois terminées
de maniére dendroide; dépdts abondants de cristaux.
Spores fusiformes, 18-21 x 7-9-(10) um, au contenu gras, 4 paroi
non cyanophile ni amylolde.
477
Récolte examinée: ZAIRE: Mayidi, leg. H. Yanderyst, ler février 1910,
HOLOTYPE, (in BR).
Nous n'avons pas retrouvé cette espéce en Afrique. Elle est proche
d'E. macarangae qui différe surtout par l'absence d'hyphes squeletti-
ques. L'E. interrupta sensu Boquiren est peut-étre E. efibulata
~
(voir & cette espéce).
5. Epithele macarangae Boidin et Lanquetin sp. nov.
Pig 4 ce
Jacens, membranacea, ex alba luteola, e parvis tuberculis, sparsis,
brevibus, concoloribus, faciliter caducis, cicatrices orliculares reli-
quentibus, constans. Super stratum hypharum horizontalium, x 2,5- 3 un,
fibulatarum, tunica tenui vel incrassata, adsunt hyphae subhymeniales
ad perpendiculum erectae, distinctae. Emergentiis 50 x 50 um, ex
hyphis tunica tenui, superne saepe dendroideis, constantibus. Basidiis
30-45 x 10-12 um, inferne subito angustatis, 4 grandibus sterigmatis.
Sporis ellipsoideis, 14-18 x 6,5-8,2 um, tunica tenui, haud amyloidea
neque cyanophila, adipabus abundantibus. In ligno demortuo. Holotypus
LY 5395. Centrafrique.
A la récolte, @étalé, poruleux puis continu, membraneux, détachable
par fragments, blanc un peu jaunatre (5 Y 9/3 a 8/4), avec marge nulle
ou fibrilleuse; émergences éparses, courtes.
En herbier, membranuleux mince, densément fendillé, créme (2,5 Y
8,5/4),orné de petits tubercules peu serrés, courts, concolores; fra-
giles et facilement caduques, ils laissent une cicatrice circulaire.
En coupe, épais de 75-100 um; il est formé d'une couche d'hyphes
horizontales x 2,5-3 um, bouclées, 4 paroi mince 4 un peu épaissie
prenant avec 1'age l'aspect d'une couche trés dense aux hyphes moins
distinctes 4 paroi souvent é@paissie (la paroi peut atteindre 0,8 um),
mais ramifiées et bouclées. Cette couche basale est haute de 20-50 um.
Au-dessus se redressent les hyphes sous-hyméniales distinctes, x2-3 um,
bouclées 4 paroi submince portant de grosses basidioles serrées, tra-
pues, largement claviformes puis subcylindriques parfois un peu étran-
glées, entre lesquelles émergent de nombreuses hyphes dendroides 4
stipe x 2 um, a paroi mince.
Basides 30-45 x 10-12 um au sommet, 4 4 forts stérigmates longs de
10-12 um; leur base bouclée est trés fortement rétrécie, parfois avec
un court stipe hyphifrome (podobaside).
Les @mergences caduques, sont généralement absentes des coupes;
larges de 50 um, elles émergent aussi d'environ 50 um et sont formées
d'hyphes 4 paroi mince x 3-3,5 um, souvent terminées en dendrophyses
complexes; elles peuvent porter des dépdts cristallins.
Spores un peu créme en masse, ellipsoides de face, ellipsoides 4
subcylindriques de profil et un peu déprimées sous l'apicule, 14-18-
(20) x 6,5-8,2 um (X = 16,68 + 1,50 x 7,42 Bs 0,42 pour le type); leur
paroi est mince, non amyloide, ni cyanophile, leur contenu riche en
graisses.
479
Récoltes: REPUBLIQUE CENTRAFRICAINE: LY 5395, HOLOTYPE, sur branchet-
tes mortes, forét touchant la savane de Bébé prés Boukoko,
1] mai 1945; LY 5410, sur Macaranga spinosa (Euphorbiacée),
La Maboké, 12 mai 1965; LY 5478, sur branchette suspendue,
Boukoko, 18 mai 1965.
Cette espéce est facile 4 reconnaitre.de la plupart des Epithele
décrits ici 4 ses émergences éparses, obtuses, fragiles et méme ca-
duques laissant une cicatrice circulaire, et l'on pourrait hésiter 4
la placer dans le genre Epithele; Toutefois ses basides et dendrophy-
ses sont du type habituel dans ce genre. Elle est proche d'E. interru-
pta Bres. par ses émergences et ses spores, mais ne montre aucune
hyphe squelettique dans son contexte.
Caractéres culturaux d'E. macarangae
SPORES.- uninucléées (LY 5395).
Douze jours aprés la dispersion des spores, de trés rares germina-
tions Sent apparues. Observées un mois plus tard les 3 germinations
isolées sont formées d'hyphes 4 cloisons simples, alors qu'a cette mé-
medate la culture polysperme montrait des hyphes étroites, réguliéres,
x 1,5-3 4, a boucles constantes. L'esp&éce est donc vrésumé hétérothalle.
Malheureusement stockées a 4°C, ces cultures ont péri avant d'avoir
pu €tre @étudiées plus complétement.
6. Epithele malaiensis Boidin et Lanquetin sp. nov.
Fig. 4 B
Tenuibus maculis, bene limitatis, maxime adhaerentibus, ex albis
griseis vel pallide alutaceis formata, 30-40 um crassa inter emergen-
tilas conicas, 200-250 x 35-40 um, quae ex hyphis skeleticis rigidis
constant. Hyphis genetricibus fibulatis; inter basidia 30-45 um longas,
tetrasporas, adsunt nonnullae hyphae dendroideae. Sporis oblongis e
facie visis, 17-21 x 8-10 um, tunica paulum incrassata, haud amyloi-
deis neque cyanophilis. In Palmaceis. Holotypus: LY 8252, Singapour.
A la récolte, en taches bien délimitées, minces, trés adhérentes,
blanc grisatre (10 YR 8/1) atteignant gris alutacé pale (10 YR 8/2,5)
au centre, 4a marge brusquement amincie; il est couvert dés la marge
d'émergences concolores, réguliéres.
Figure 4: A- Epithele nikau Cunn.: 1- coupe du basidiome au niveau
d'une émergence, et spores (LY 9119), in NH4OH 60°C-
Congo; 2- spores de L'holotype (in K).
B- Epithele malaiensis sp. nov.: spores de lL'holotype
CLY* 8252).
C- Epithele macarangae sp. nov.: spores de l'holotype
(LY $395).
480
En herbier, trés adhérent, blanc a4 alutacé pale (10 YR 8/3 a 8,5/4).
En coupe, il apparait trés mince (30-40 um) entre les émergences
~
c6niques de longueur totale de 200-250 um, larges de 35-40 um 4 la
sortie de l'hyménium, plus larges 4 l'enracinement qui se fait sur le
substratum.
Emergences constituées d'hyphes squelettiques rigides, x 1,8-2,5 um
sensiblement congophiles aprés traitement ammoniacal 4 60°C qui dissout
un dépét plus ou moins cristallisé qui les englue; a 1'écrasement
quelques hyphes dendroides s'en détachent et l'on voit a la base que
les fibres sont un peu plus larges 4 paroi plus é@épaissie et sont
Sinueuses. Entre les émergences quelques hyphes génératrices peu dis-
tinctes, bouclées, portent des hyphes sous-hyméniales 4 paroi submince;
hyménium formé de grosses basidioles ovoides puis claviformes 4 sub-
cylindriques, riches en gouttelettes réfringentes, et d'hyphes den-
droides gréles; basides fortes, longues de 30-45 um brusquement rétré-
cies 4 la base qui est bouclée, portant 4 gros stérigmates; leur lar-
geur atteint 15 um a4 mi-hauteur.
Spores oblongues de face, elliptique de profil 17-21 x 8-10 um
(x = «18,45 * 0,80 x 8,82 % 0,50), 4 paroi trés peu épaissie (0,3 290,5
um dans KOH 3%) non cyanophile, ni amyloide, au contenu gras.
Récoltes : SINGAPOUR: LY 8252 HOLOTYPE, sur Paltiae, Mac Ritchie
reservoir, 19 aoiit 1977.
A les plus grandes spores parmi les Epithele 4 émergences squelettiques
Caractéres culturaux d'E. malaiensis
SPORES.- uninucléées (LY 8252)
POLYSPERME.—
Croissance: extrémement rapide (botites couvertes en une semaine)
Aspect: marge réguliére . Mycélium aérien, blanc peu développé dans
la partie jeune ot il est réduit 4 une grosse pruine discontinue puis
forme un feutre trés mince, régulier, sur lequel sont dispersées des
pustules coriaces d'aspect céracé puis recouvertes d'une couche cray-
euse formant souvent bourrelet circulaire quand elle n'atteint pas le
sommet au centre de la pustule. Sous le couvercle, mycélium blanc 4
aspect rhizoide. Observée par transparence, la culture montre en outre
quelques lignes blanches plus denses. Le revers des cultures est
inchangé, l'odeur nulle.
Microscopie:
Mycélium aérien: hyphes gréles, x 0,8-1,2 wm, bouclées,a paroi min-
ce, contenu homogéne,cachant des hyphes, x 1,8-4,5 u, bouclées.
- partie hyaline coriace, céracée, des pustules: hyphes ramifiées,
densément enchevétrées mais non soudées, x 2,5-3 um, bouclées, 4 paroi
réfringente un peu épaissie mais plus souvent au contenu dense, homo-
géne, réfringent.
- partierecouvrante blanche: hyphes x 2-2,5 um 4 paroi submince ou
parfois nettement épaissie, bouclées non soudées, mais trés intriquées
en sorte de puzzle.
Mycélium submergé: hyphes, x 1,5-5-(6) um, 4 paroi mince, contenu
homogéne et boucles constantes. Toutefois quelques hyphes bouclées 4
paroi nettement et irréguliérement épaissie ont été rencontrées.
Les lignes blanches s'enfongant dans la gélose sont constituées
d'hyphes, x 2-2,5 um, trés intriquées en sorte de puzzle.
Boucles: constantes.
481
Cytologie: hyphes aux articles binucléés.
Oxydases: ac. gallique: +++,0 ga1acolL: +++, 0
p.- crésol: -, précipité blanc tyrosine: =, .0
CODiemeza- 36-= (11) = 32 -736,-7390,-74)" = 54. = 64,
Remarque. Cette culture de croissance extrémement rapide est caracté-
risée par la formation de pustules céracées.
7. Epithele nikau G.H. Cunn.,Trans. Roy. Soc. New Zeal.
B22 029,1 11956,
Fig. 4A
A la récolte, en petites taches séparées puis largement confluent,
trés étalé, blanc grisatre (10 YR 7,75/1) A bordure blanche étroite;
émergences nombreuses, concolores.
En herbier, adhérent, blanc plus ou moins créme (5 Y 8,5/3) couvert
de petits aiguillons concolores nombreux jusqu'a la marge qui est trés
étroite, atténuée, blanchatre.
En coupe, @épais de 30-50-80 um hors des émergences qui mesurent 180-
230 x 12-25 um et dépassent l'hyménium de 70-100-190 ym. Ces émergences
sont cOniques dans leur partie inférieure puis subcylindriques et sont
formées d'hyphes rigides a paroi épaissie, x 1,8-2,2 um, un peu rétré-
cles au sommet, assez congophiles, en faisceau régulier, le plus sou-
vent enrobées d'une substance hyaline réfringente, cassante, non biré-
fringente qui englue les hyphes externes du faisceau. A la base 1'émer-
gence atteint le substratum en s'élargissant; elle est alors formée
d'hyphes fibroides congophiles maissinueuses ramifiées rappelant par-
fois des binding-hyphae (cf. Corner 1953).
La base est formée sur 15-18 um d'hyphes génératrices horizontales,
x 1,8-2 um, réguliéres, 4 paroi mince, bouclées, auxquelles se mélent
hors des émergences de rares éléments squelettiques, qui se multiplient
aux abords de celles-ci avant de s'emméler entre elles pour former
leur base.
Hyménium haut de 20-35 um, formé de basidioles et basides riches en
huiles; les basides ou podobasides larges, légérement étranglées aux
2/3 supérieurS,mesurent 18-27 x 6-8 um au sommet, et sont plus larges
au 1/3 inférieur (x 7,5-9,5 um). Elles portent 4 forts stérigmates
souvent arqués. Quelques hyphes simples ou faiblement dendroides émer-
gent entre les basidioles riches en gouttes grasses.
Spores ellipsoides de face, fusiformes de profil, 10-14 x 4,5-6 Um
a paroi mince non amyloide, ni cyanophile, au contenu guttulé; l'api-
cule est trés réfringent; x = 12,16 = 1,10 x 5,52 * 0,35 pour 8004
12,4397. 0, 709x501, 20,27. pours9l 1g
oe
Récoltes: GABON: LY 8004, sur Ancistrophyllum secundiflorum,
La Mondah, 30 mai 1976; LY 9119, sur Eremospatha macrocarpa,
id, 20. janvier: 1978, lee. G. Gilles 1406.
482
Caractéres culturaux d'E. nikau
SPORES.- (LY 8004) uninucléées
GERMINATIONS.- Apparues quatre jours aprés la dispersion des spores
LY 9119, elles ont permis 25 isolements.
MONOSPERMES.- A sept jours, les cultures montrent une majorité d'hy-
phes 4 cloisons simples, mais aussi de rares petiteshyphes bouclées.
A un mois la moitié des cultures sont totalement bouclées et 4a 2
mois, toutes les cultures sont binucléées, 4 boucles constantes.
L'espéce est donc présumée homothalle.
POLYSPERME.- (LY 8004, LY 9119)
Croissance: rapide (boites couvertes en trois semaines)
Aspect: la marge est réguliére. Mycélium aérien blanc pur, appliqué,
peu abondant, ne cachant pas totalement le milieu. Plus dense, feutré-
ruguleux vers la bouture, il est partout ailleurs un peu rayonnant
(LY 9119) Aa nettement rayonnant chez LY 8004 ot il forme d'étroites
fibrilles disposées trés réguliérement et dans la partie jeune des
cultures, ces fibrilles, serrées, paralléles,grimpent contre le bord
des boites pour s'étaler ensuite sous le couvercle. Le revers des
cultures est inchangé, leur odeur nulle.
Microscopie:
Mycélium aérien: hyphes, x 1-2,5-(4,5) wm, réguliéres, a paroi min-
ce, boucles constantes. Les hyphes gréles, x 1-1,2, ont souvent des
boucles un peu fortes. Dans le mycélium jeune, quelques hyphes sont
incrustées; dans le mycélium agé les dép6ts de cristaux biréfringents
sont plus importants.
Mycélium submergé: hyphes bouclées, les unes trés gréles, x 1-1,2 um
trés réguliéres, les autres plus larges, irréguliéres, x 2,5-4-6 um, 4a
parol mince,a contenu hétérogéne. Celles-ci surtout nombreuses 4 fai-
ble profondeur montrent quelques renflements atteignant 10 ywm.On peut
aussi observer de rares hyphes 4 paroi, x 0,7 um, également bouclées.
Boucles: constantes
Cytologie: hyphes aux articles réguliérement binuciéés, bouclés.
Oxydases: ac. gallique: +++++, 0 gaiacol: +++++," 0
p.= crésol: ~-, fort précipité blanc tyrosine?) =," tread
COUR MH 2a = SCD eV 2h ash sO 4 oie ae) ee od 5 mm
8. Epithele ovalispora Boidin et Lanquetin sp. nov.
Fiz,. 5
Jacens, membranacea, alba, margine paulum sub lente fibrillosa, acu-
bus exilibus hirta, 100-400 um crassa. Emergentiis 150-280 x 30-35 um,
ex hyphis tunica incrassata, rigidis, tenui strato hypharum fibulata-
rum, tunica tenui circumdatis constantibus. Trama ex hyphis genetrici-
bus fibulatis. Basidiis 45-55 x 10-13 um, 4 grandibus sterigmatis.
Sporis ovatis, 13,5-17,5 x 8,5-11 um, tunica haud amyloidea neque
cyanophila. In Palmaceis. Holotypus LY 7928. Gabon.
A la récolte, étalé, membraneux mais adhérent, 4 marge un peu fibril-
leuse radiée sous la loupe, hispide par des aiguillons nombreux, longs
et effilés, blanc.
484
En herbier, il a le méme aspect mais il est teinté de créme (vers
2,5 Y 9/3) sauf 4 la marge blanche et sans émergence sur Q,5 mm
environ.
En coupe, épais de 100-140 im dont les 30-40 um inférieurs forment
une couche d'hyphes basales horizontales: ce sont des génératrices 4
paroi mince ou submince, bouclées, x 2-3-(3,8) um. Au sommet de la cou-
che inférieure naissent d'une part les émergences, d'autre part les
hyphes sous-hyméniales assez laches, x 2,5-5 um bouclées.
Les émergences sont céniques, longues de 150-230-280 um, étroites,
elles débordent de 70-160-(230) um et sont larges de 30-35 um 4a la
sortie de l'hyménium; elles sont constituées d'hyphes, x 2 um environ,
d'aspect fibreux, congophiles et assez dextrinoides et cyanophiles, 4
paroi assez épaissie, serrées parallélement en faisceau dans la par-
tie émergente; a la surface des émergences ont peut voir une mince
coucke d'hyphes génératrices bouclées.
Basidioles largement claviformes, larges de 10-14 wm, riches en
huile. Certaines (en arrét de végétation?) ont une paroi épaissie.
Entre les basidioles 6émergent des hyphes dendroides gréles, x 2 um
environ. Basides larges, 45-55 x 10-13 um au sommet, plus larges plus
bas: Gxul4-15 um), a 45 forts: stérigmates 15-16 x-3,5-—4. um.
Spores ovoides de face, au contenu gras,013, 5-1 7,5 x 835-11) im
(x = 15,05 = 1,12 x 9,70 + 0,65) a paroi non amyloide, ni cyanophile;
en masse, la sporée est teintée d'orange brunatre.
Récoltes: GABON: LY 7928 HOLOTYPE, sur Ancistrophyllum secundiflorun,
rive gauche de l'Ivindo, Makokou, le 24 mai 1976.
Caractérisé par des émergences longues et cdniques, insérées
a mi-hauteur dans le champignon et aux hyphes congophiles et cyano-
philes , et sutout, par ses spores.
9. Epithele sp. LY 5697 d'Argentine
Fig. 6A
En herbier, membraneux, blanc créme, détachable, comme posé sur un
subiculum aranéeux lachement feutré formant une étroite marge amincie
blanche; couvert sous la loupe d'aiguillons gréles, nombreux.
En coupe épais de 200-300 um avec couche inférieure lache trés
développée supportant un hyménium dense, épais de 50 um environ.
Emergences 70-120 x 30-50 um, formées d'hyphes x 2-3 um bouclées, 4
paroi mince ou un peu épaissie parfois ramifiiées, souvent terminées
en éléments dendroides; dans l'axe de ces émergences on peut voir quel-
ques hyphes 4 paroi épaisse, certaines sont des hyphes bouclées sur
leur parcours, d'autres peu nombreuses sont des extrémités fibroides.
Ces émergences portent des dépdts de cristaux.
La couche inférieure est formée d'hyphes trés distinctes, les généra-
trices bouclées, larges de 2-5 um, a parol mince ou un peu épaissie,
mais facilement collapsées, hyphes squelettiques réguliéres, x 3-3,5 um
a lumen subnul.
Basides claviformes pui
stérigmates.
Spores ellipsoides, a i
mince ou faiblement épaissie, non amyloide, ni cyanophile.
s subcylindriques, 28-37 x 6-7 um, a 4 gros
10-12 x 5-6,5 um, avec apicule brillant, a paroi
485
Figure 6: A- Epithele sp.5697 d'Argentine:baside,sporés vues sur le
basidiome et hyphes du contexte(Congo ammoniacal).
B- Pteridomyces lacteus sp.nov. :sommet d'une émergence,
spores(holotype LY 9334)et basides;a-arthrospores en culture.
C- Vesiculomyces epitheloides: sulfocystides et sorte
d'arthrospores en culture (LY 96/77,paratype).
486
Récolte: ARGENTINE, sur bois sec, El Sgherbio, Misiones, 28 avril
1966, leg. C.E. Gomez.
Cette espéce a été signal@ici car elle différe assez nettement
des Epithele décrits ci-dessus. Son contexte lache avec hyphes squelet-
tiques dominantes, ses émergences pourvues, au contraire,de quelques
hyphes squelettiques enrobées d'hyphes bouclées qui ne semblent pas
s‘enraciner profondément, en font une espéce marginale bien typée;
elle mériterait d'étre décrite, mais a pu 1'étre dans un autre genre
(Hydnum peut+vétre) par d'anciens auteurs latino-américains... ce qui
nécessiterait une révision de nombreux types difficilement localisables.
II - Genre Pteridomyces Julich 1979 emend.
Champignons étalés, 4 marge adhérente, ornés d'émergences stériles
granuliformes ou odontioides nées soit en surface, soit plus ou moins
profondément; contexte et émergences monomitiques ou dimitiques. Den-
drophyses ou hyphes dendroides. Basides claviformes petites 15-20-(30)
x 4-5 um, a (2)-4 stérigmates. Spores petites (moins de 9 um de lon-
gueur), A parol mince, ni amyloide, ni cyanophile.
10. Pteridomyces lacteus Boidin, Lanquetin et Gilles sp. nov.
Fig. 6 B
Orbicularis, deinde confluens, late expansus, adhaerens, e cretaceo
albus vel pallidissime cremeus, 40-100 um crassus. Emergentiis cylin-
dratis, brevibus, 50-75 x 20 um eminentibus. Trama monomitica, fibu-
lata. Basidiis 22-30 x 5 um, (3)-4 sterigmatis exilibus 7-8 um longis,
nonnulis hyphis dendroideis intermixtis. Sporis ellipsoideis, 6-8 x
3-4 um, tunica tenui, haud amyloidea neque cyanophila. In ligno.
Holotypus LY 9334. Gabon.
A la récolte, orbiculaire puis confluent,blanc de craie ou 4 peine
jaunatre (2,5 Y 8/0,5) avec marge amincie.
Emergences courtes,cylindriques larges de 15-20 um.
En herbier, largement étalé, mince trés adhérent, blanc 4 créme trés
pale, a marge nette, amincie sur 90,2-0,4 mm, blanche. Emergences petites,
visibles sous la loupe comme de minuscules aiguillons concolores régulié-
rement répartis sauf vers la marge.
En coupe, @pais de 40-100 um avec émergences atteignant 100 um de
longueur totale, débordant de 50-75 x 12-20 um. Monomitique: une
couche basale haute de 10-20-(30) um est formée d'hyphes horizontales
a parol mince ou submince, réguliéres, bouclées, x 2,5-3,8 um; se
dressent ensuite un sous-hyménium fait d'hyphes semblables plus rami-
fiées et un hyménium fait de baidioles claviformes étroites et de
petites basides 22-30 x 5 um, 4 3-(4) stérigmates longs de 7-8 um.
Entre les basidioles peuvent émerger des hyphes dendrophysoides. Les
émergences sont faites d'hyphes 4 paroi mince, ou a peine épaissie
dans la partie immerse, un peu sinueéuses, bouclées, x 2-3 um.
487
Spores ellipsoides inégalement attenuées (de profil) vers l'apicule,
6-8 x 3-4 ym, a4 paroi non amyloide. (x = 7,01 x 3,22 pour le type ,
i200 pour LY -9338)".
Récoltes: GABON: LY 9334, HOLOTYPE, sur bois mort tombé de dicoty-
lédone, fourré littoral 4 14 km au N. de Libbreville,
Simai 1979, leg. Gi Giltes 15745 LY-9338, abid., leg:
G. Gilles 1576.
Caractéres culturaux de Pteridomyces lacteus
SPORES.- uninucléées (LY 9334, HOLOTYPE)
GERMINATIONS.- (LY 9334 et 9338). Elles apparaissent 3 ou 4 jours
aprés la dispersion, 30 isolements 9334 et 8 de LY 9338 ont été
effectués:
MONOSPERMES.- Ils sont formés d'hyphes aux articles uninucléés, sans
boucles. L'appariement des monospermes 9334 n'a pas permis de définir
le type de thallie avec certitude. Il se dégage une tendance tétra-
polaire. Les monospermes LY 9334 sont totalement intercompatibles
avec les monospermes LY 9338.
POLYSPERME.— LY 9334.
Croissance: trés rapide (boites couvertes en deux semaines)
Aspect: La marge est trés réguliére. A 6 semaines, sur un milieu
irréguliérement teinté de bai (2,5 YR 4/6) ou de chatain (2,5 YR 3/4),
on observe un mycélium subfeutré, velouté, blanc 4 créme rosé (5 YR
8/3) de densité variable, beaucoup plus faible sur les zones de
milieu brun rougeatre qu'il ne cache pas totalement. Dans d'autres
cultures, on observe deux parties trés contrastées: une grande plage
de mycélium subfeutré blanc sur milieu non teinté et une grande zone,
englobant la bouture, ot le milieu bruni est recouvert d'un mycélium
alutacé (10 YR 8/4 a4 8 YR 7/6). Le revers des cultures est teinté de
brun roux par plages, l'odeur est nulle.
Microscopie:
Mycélium aérien: dans les parties blanches, il forme une peau consti-
tuée de nombreuses hyphes superficielles trés gréles (qui sont en fait
des hyphes collapsées) cachant des hyphes x 2-4um, réguliéres, 4
paroi mince et boucles constantes. Dans les jeunes cultures, certains
rameaux se désarticulent en arthrospores (cf. fig. 6 B a).
Dans les parties teintées, le mycélium superficiel coloré est formé
d'hyphes bouclées, parfois x 1-1,5 um mais jusqu'a 8-9 Um. Certaines,
x 1,8-4 ym, ont une paroi épaissie comme rigide, trés réfringente, jau-
ne orangé, elles portent des gouttes résinoides de méme couleur ou
sont remplies de cette substance. Elles peuvent former des amas denses
brun doré sous le microscope. Aprés trois mois, les zones teintées
montrent sous une mince couche d'hyphes affaissées, une crofite coriace,
impossible a dilacérer; elle se casse montrant sur les bords des
hyphes irréguliéres, trés intriquées, 4 paroi épaisse, réfringente
et aucontenu jaune doré.
Mycélium submergé: hyphes, x 2-5-(8) um, souvent larges, réguliéres
ou parfois irréguliéres (difformes, renflées), 4 paroi mince et boucles
constantes. Localement des masses compactes sont formées d"hyphes
bouclées, serrées, intriquées,x 3 ym environ, hyalines 4 brun doré,
a paroi épaisse 4 trés épaisse plus ou moins gélifiée.
Boucles: constantes.
Cytologie: hyphes aux articles binucléés et bouclés.
488
Qxydases: ac. gallique: +++++, O galacol:t+t++. 0
p-— crésol: -, précipité blanc tyrosine: -, 5 mm
CODE: 2a = 8c) =035. = 3/7) = 39:— .42 -954 = 58) =" 6)
Remarque. Cette culture est caractérisée par ses plages brun-rou-
geatres, la formation d'arthrospores, et la présence d'amas denses
en mycélium submergé.
1] - Pteridomyces roseolus Boidinet Lanquetin sp. nov.
Deters Fig. /
Erumpens, jacens, subceraceus firmus, e roseo griseus, permultis
emergentiis gracillimis coopertus, 35-50 um crassus. Emergentiis
50-65 x 15-20 um transgredientibus, in medio ex hyphis tunica incras-
sata et extus ex hyphis fibulatis, tunica parum incrassata constanti-
bus. Hyphis basalibus subhymenialibusque fibulatis, tunica tenui.
Basidiis 18-20 x 3,8-5. um. Spords subcylindratis, 5=6.x 2-2, 5m,
tunica haud amyloidea neque cyanophila. In ligno. Holotypus 5377.
Centrafrique.
A la récolte, é@rompant, étalé, mince, subcéracé ferme, gris rosé
(5 YR 6/2, 6/3) couvert sous une forte loupe de multiples émergences
gréles, hyalines.
En herbier, aride, beige pale (vers 10 YR 7,5/2) a émergences pales,
serrées, gréles, a peine visibles sous la loupe.
En coupe, @pais de 35-50 um entre les émergences posées sur le sup-
port et mesurant dans leur totalité 90-100 um de longueur; elles émer-
gent donc de 50-65 um; leur forme est un peu cdnique, leur largeur est
de 15-30 Um au niveau des basides.
Les émergences sont solides, formées d'hyphes cohérentes trés dif-
ficiles a écarter; elles sont formées en surface d'hyphes x 2 unm,
bouclées, parfois ramifiées, 4 paroi un peu épaissie; les hyphes
plus profondes ont une paroi plus épaisse et nacrée. Aprés traitement
ammoniacal a 60°C les hyphes des émergences apparaissent trés sinueu-
ses a labase, larges de 1,5-2 um, a paroi trés épaissie (gonflée?),
trés congophile et 4 lumen indiscernable; si on les place dans le
bleu aprés ce traitement ammoniacal 4 chaud, le lumen bleuté n'est net
que sur les hyphes de surface, les autres en semblent dépourvues.
Entre les émergences on voit des hyphes gréles x 1,5-2 ym, serrées,
plus ou moins ascendantes, 4 paroi mince ou un peu épaissie, bouclées,
formant une couche basale localement bourrée d'amas de cristaux, et
une zone hyméniale constituée de petites basidioles claviformes bou-
clées et de quelques hyphes émergentes non dendroides.
Basides 18-20 x 3,8-5 ym a4 4 stérigmates.
Spores subcylindriques, 5-6,2 x 2-2,5 wm 4 paroi mince, non amyloide,
ni cyanophile.
Récolte: REPUBLIQUE CENTRAFRICAINE: LY 5377, HOLOTYPE sur bois mort,
La Maboké, 10 mai 1965.
Il est difficile ici de se prononcer sur la nature squelettique ou
non des hyphes des émergences. Aprés traitement ammoniacal a 60°C les
hyphes congophiles et 4 paroi trés épaissie, qui constituent 1l'essen-
tiel des émergences, rappellent les hyphes "squelettiques" caractéris-
tiques des émergences de Skeletohydnum. Une observation directe 4
froid montre clairement que la surface des émergences est constituée
d'hyphes génératrices bouclées parfois ramifiées mais 1'intérieur
489
Figure 7: Pteridomyces roseolus sp.nov.: coupe du basidiome ,
avec detail d'une partie d'une émergence (Ammoniaque
60°C puis Bleu Coton); spores (holotype LY 5377).
est alors formé d'hyphes trés réfringentes, solidement accolées, que
l'on ne peut disjoindre et distinguer individuellement sur d'assez
longs parcours. Basides et spores sont du type Pteridomyces.
On aurait donc ici un Pteridomyces Aa émergences constituées d'hyphes
squelettiques ou au moins 4 nette tendance squelettique. Nous
l'avons retenu ici, bien que nous ne disposions que d'une seule
récolte, parce qu'il pose le probléme de l'utilisation du "mitisme"
dans la définition des genres épitheloides.
490
12. "Pteridomyces" sp. 8270 de Singapour
Cette récolte de Singapour n'est citée ici que parce qu'elle pose
elle aussi le probléme du mitisme chez les Pteridomyces. N'ayant pas
sporulé aprés la récolte, nous ne voulons en faire une récolte type.
En herbier, étalé, trés adhérent, mince, beige sale (10 YR 7/2,5)
a marge amincie sur un ou deux millimétres, grisatre (vers 5 YR 7/1),
couvert sous une forte loupe de petites émergences concolores ou 4
sommet bruni, serrées, @parses ou alignées.
En coupe, épais de 70-170 um entre les émergences, il est constitué
d'hyphes serrées, peu distinctes, 4 paroi souvent épaissie, bouclées,
qui portent des basidioles claviformes petites et des basides d'envi-
ron 14 x 4,5 um, a 4 stérigmates longs de 4 4 5 um.
Les @mergences, trés nombreuses, débordent de 50-70 x 15-30 ums
elles sont ancrées profondément et constituées d'hyphes squelettiques
congophiles raides, x 1,5-3 um, a lumen étroit; ces émergences sont
recouvertes de dép6ts cristallins.
Spores (vues en petit nombre) oblongues, 3,5-4 x 2,5-2,8 um, non amy-
loides, ni cyanophiles.
Récoltes: SINGAPOUR: LY 8270, sur bois, Mac Ritchie Reservoir, 19 aoit
LOR de
Cette récolte rappelle beaucoup par son aspect P. roseolus.
Par ses basides et ses spores il appartiendrait au genre Pteridomyces
mais ses émergences sont constituées d'hyphes squelettiques tout 4
fait typiques comme chez les meilleurs Skeletohydnum.
13. Pteridomyces sphaericosporus Boidin, Lanquetin et Gilles sp. nov.
Figs 8
Jacens, e griseolo albus, dein alutaceus, lentus, spongiosus, emer-
gentiis albidis hirtus, 150-180 um crassus inter emergentias, quae
60-90 x 20-30 umtransgrediuntur. Strato inferiore laxo, ex hyphis
fibulatis tunica crassa sicut.in emergentiisconstante. Basidiis 18-20
x 4 um. Sporis subglobosis, 4,5-5,6 x 3,8-4,8 um, tunica paulum crassa,
haud cyanophilis neque amyloideis. In ligno. Holotypus LY 7377.
Céte d'Ivoire.
A l'état frais, largement étalé, blanc grisatre (10 YR 8/1) mais
devenant pale alutacé (10 YR 8/3,5) dés qu'il séche; coriace,spongieux,
trés hérissé d'émergences blanchatres; marge concolore, abrupte ou
amincie.
En herbier, étalé, adhérent, alutacé (10 YR 8/4 4 7,8/4) finement
granuleux sous la loupe ot il apparait aranéeux dense plus poruleux,
enfin continu avec émergences subcylindriques un peu plus pales
apparaissant dés la marge irréguliérement groupées, parfois juxtaposées
ou connées par trois ou quatre.
492
En coupe, épais de 150-180 um, sous les émergences qui mesurent
60-90 x 20-30-(40) um; il est nettement constitué de deux parties:
une couche inférieure lache, haute de 100-120 um, formée d'hyphes en
tous sens et d'un hyménium dense épais de 60 ’m environ. La couche
inférieure est faite d'hyphes trés distinctes, réguliéres, bouclées,
a paroi trés épaisse, hyaline, larges de 2-4 um, ramifiées. Leur paroi
reste nettement €épaissie aprés les premiéres ramifications qui forment
le sous-hyménium. Sous-hyménium aux articles beaucoup plus courts,
a parol mince ou un peu épaissie, aux boucles étroites.
Basides subcylindriques, 18-20 x 4 um, a4 4 stérigmates de 2-2,5 um
de longueur. Trés souvent les éléments hyméniens, hors végétation,
contiennent des hyphes 4 1'intérieur, 4 paroi mince, souvent rata-
tinées.
Emergences constituées par des faisceaux d'hyphes x 4 um environ,
a paroi hyaline trés é@paissie (1 4 1,5 um), nées souvent trés bas,
bouclées sur leur parcours, souvent sinueuses ou étranglées et plus
ou moins moniliformes, piquetées de cristaux sur les 30 um terminaux.
_Spores subsphériques 4 un peu ovoides, 4,5-5,5 x 3,8-4,8 um,
(x = 4,98 + 0,33 x 4,20 £0,25) 4 paroi un peu épaisse, non amyloide,
ni cyanophile, contenant un globule gras.
Récolte: COTE D'IVOIRE, LY 7377, HOLOTYPE sur bois, en forét du Téké,
7) juritet® 19745 “lee. CG. Gilles 248.
Facile a reconnaitre a ses spores et a sa structure, ce qui en fait
un membre assez marginal du genre Pteridomyces.
~
III - Un champignon épitheloide 4 spores amyloides
Il nous reste a décrire un champignon malgache qui n'entre pas dans
les genres @étudiés ci-dessus. Il posséde, @mergences mises 4 part, les
caractéres du genre Gloeocystidiellum Donk.
Ce genre était certes commode mais hétérogéne comme 1l'ont signalé
divers auteurs tel Boidin (1966) dans un tableau hypothétique (pl. II)
des relations des espéces de ce genre avec diverses familles. Depuis,
des coupures successives ont été proposées: genres Scytinostromella
Parmasto (1968), Vesiculomyces Hagstrom (1977), Megalocystidium Julich
(1978), Conforticium Hallenberg (1980), Boidinia Stalpers et Hjortstam
(1982). Il semble que le genre Megalocystidium comprenne les espéces
a spores lisses -ce qui n'est pas dit dans la diagnose originale, qui
ne signale aucun caractére distinctif par rapport a Gloeocystidiellum
porosum type du genre démembré, mais qui a, lui, des spores ornées.-—
En effet le type de ce nouveau genre, M. leucoxanthum et les deux
autres espéces transférées par Julich, M. lactescens et luridum, ont
des spores lisses. Mais c'est aussi le cas du genre Vesiculomyces
Hagstrom proposé antérieurement. Son auteur le caractérise avant tout
par ses gloeocystides sulfoaldéhyde négatives. V. citrinus posséde
en fait des gloeocystides trés précocement vides dans le carpophore,
mais qui, en culture, ont un contenu sulfo-positif (Boidin 1958 p. 109).
493
L'absence de boucles, autre caractére proposé comme distinctif se re-
trouve chez G. lactescens (un Megalocystidium pour Julich) et G.
humile. Nous ne voyons donc aucune raison pour ne pas synonymiser
Megalocystidium a Vesiculomyces antérieur d'une année.
Genre Vesiculomyces Ha Hagstrom (Bot. Nat. 130: 53, 1977) emend.
Ce genre est réservé aux Gloeocystidiellum 1 monomitiques 4 spores
amyloides lisses, pourvus de gloeocystides montrant le plus souvent
un contenu riche, pouvant réagir aux sulfo-aldéhydes, contenu qui
parfois cependant disparait précocement dans le basidiome. Les hyphes
sont avec ou sans boucles. Contrairement au genre Conferticium
Hallenberg, ici les hyphes ne sont pas cyanophiles et les basides ne
~
repoussent pas 4 l'intérieur de basides anciennes (répétobasides).
Le type est Vesiculomyces citrinus (Pers.) Hagstrom.
Autres espéces: Vesiculomyces Chelidonium (Pat. Bull. Soc. Mycol.
France, 16: 180, 1900, ut Corticium) Boidin et Lanq. nov. comb.,
V. humilis (Boidin, Cah. Maboké, 4: 9, 1966, ut Gloeocystidiellum)
nov. comb.; V. leucoxanthus (Bres. Fungi Trid. 2: 57, 1898, ut
Corticium) nov. comb.; V. lactescens (Berk. ap. Smith Engl. Fl. 5,2:
169, 1836, ut Thelephora) nov. comb.; V. luridus (Bres. Fungi Trid.
22559, 1898, ut Corticium) nov. combs; V. sulcatus (Rehilil et Bakshi,
Indian For. Bull. 242: 15, 1965, ut. Corticium) nov. comb.
- Vesiculomyces epitheloides Boidin, Lanquetin et Gilles sp. nov.
Fig. 9
Jacens, amplus, solidus, rigescens, salmoneus, ora alba fibrillosa,
300-350 um crassus. Emergentiis granuliformibus 60-70 x 20-60 um
eminentibus, crassamentis crystallinis uberibus. Hyphis jacentibus,
angustis, simplicibus septis, stratum ad 50 um crassum constituenti-
bus; hyphis subhymenialibus angustis, resurgentibus tunica incrassata;
hymenio increscente e gloeocystidiis obtusis, 150-250 x 10-20 um,
sulfoaldehydibus nigrescentibus, atque basidiis 45-60 x 6-8 um, tetra-
sporis constante. Emergentiis ex hyphis parallelis, x 2-3,25 um, tuni-
ca subtenui, vel paulum incrassata constantibus, sporis reniformibus,
6,>-6 * 4,5-5,)5, 1m. Holotypus LY 9761, Madagasear.
A lia rvecolte, Gtalé, étendu, solide, saumoné (7,5 YR 7/4 a 8/6,
salmon buff R.) avec bordure fibrilleuse blanche pouvant atteindre
3 mm de largeur; sous la loupe il est constellé de petites émergences
concolores.
En herbier, trés adhérent, assez corné, uniformément isabelle (7,5
YR 6,8/4,5), avec marge amincie, ocre alutacé (10 YR 8/6, warm buff),
puis fibrillo aranéeuse; sous la loupe trés nombreuses émergences
assez réguliérement disséminées, petites, granuliformes, concolores.
Coupe haute de 300-350 um sans les émergences qui dépassent de 60-
70 um et sont larges de 20-60 yum, brillantes par les amas de gros
cristaux qu'elles portent. Sur 50 ym environ d'hyphes horizontales,
étroites, x 1,5-2,5-(3) wm a paroi mince ou un peu épaissie 4 cloisons
simples, serréeS mais non soudées entre elles, une zone de transition
formée d'hyphes obliques similaires 4 paroi souvent plus nettement
épaissie, porte un hyménium crassescent haut de 180-250 um formé
d'énormes gloeocystides obtuses, irréguliérement subcylindriques,
150-250 x 10-20 ym, au contenu réfringent sulfoaldéhyde positif, 4
paroi épaissie dans la moitié inférieure, pouvant montrer quelques
10pm
spores \¥
|
| KU\_x
Figure 9: Vesiculomyces epitheloides sp.nov.:base du basidiome ;
catahyménium (X signale un raccord) ,émergence et spores
(holotype LY 9761).Coupe montée dans le Bleu lactique.
495
cloisons de retrait, d'hyphes gréles x 2-3 ym 4 paroi mince pouvant
émerger entre les basidioles, et .de basides étroitement claviformes,
a paroi mince, sans boucle 4 la base, 45-60 x 6-8 ym 4 4 stérigmates.
Les émergences sont formées d'hyphes paralléles, larges de 2-3, 25
um, a& paroi submince mais apparaissant épaissie (0,8 pm environ) aprés
traitement ammoniacal a 60°C, ramifiées, parfois un peu élargies au
sommet, mélées 4 des amas de gros cristaux. Les émergences "s'enfon-
cent" dans 1l'hyménium jusqu'a 200 ym sous la surface. ig
Spores blanches en masse, réniformes, 6,5-8 x 4,5-5,5 wm, (x = 7,49
* 0,61 x 4,95 + 0,23), 4 paroi lisse, mince, amyloide, mais non cyano-
phile et au contenu guttulé gras.
Récolte: MADAGASCAR: LY 9677, sur une souche sous un buisson, alt.
1250 m, la, Mandraka,, .29 mars I981,. les. G. Gilles n° 28;
bYoO7Ol, wabid. Ii-aveil 198.5 lee, CG. Gilles n 46,
HOLOTYPE.
Caractéres culturaux de Vesiculomyces epitheloides
SPORES.- (LY 9677, 9761: Holotype), binucléées
MONOSPERMES.- Trés peu de spores ont germé. Deux germinations 9761
et 6 germinations 9677 ont été isolées respectivement 8 et 15 jours
aprés la dispersion des spores. Ces cultures monospermes sont formées
d'hyphes réguliéres, sans boucles, aux articles uninucléés. Elles
montrent des éléments de forme irréguliére au contenu densément guttu-
1é, sulfo-aldéhyde positif et des hyphes étroites qui se désarticulent
en arthrospores.
Tests d'intercompatibilité: toutes les confrontations de monospermes
9761 avec 9677, produisent un mycélium formé d'hyphes aux articles
réguliérement binucléés. L'espéce est donc hétérothalle et les deux
récoltes 9761 et 9677 sont intercompatibles (cf. Lanquetin 1973).
POLYSPERMES. —
Croissance: extrémement lente (ne couvre pas la moitié de la boite en
six semaines)
Aspect: Marge irréguliére, plus ou moins lobée puis mycélium aérien,
blanc léger puis floconneux(pouvant aussi former des bourrelets con-
centriques) formant dans la partie agée 1 couche lisse ou bosselée-
grumeleuse, de mycélium blanc 4 ocre alutacé pale (7,5 YR 7,5 a 8/4,
a 10 YR 8/4) ou alutacé. chamois (10 YR 8/6 a 7,5/6) sur les reliefs.
Le revers des cultures est inchangé; leur odeur faible, agréable,
constante.
Microscopie:
Mycélium aérien: il montre
- des hyphes, x 1,2-3-(4) um, réguliéres, a paroi mince, aux cloisons
simples
- de nombreuses gloeocystides de tailles et formes trés variées, avec
renflements: x 2,5-5-(10) um (cf. fig. 6 C). Leur contenu réfringent,
densément guttulé ou pailleté dans le rouge congo, réagit trés posi-
tivement dans les sulfo-aldéhydes. Aucune schizopapille n'a été obser-
vée
- quelques hyphes avec cloisons de retrait qui se désarticulent en
arthrospores (cf. dessin)
496
Mycélium submergé: dans le premier millimétre de gélose un peu coriace
a couper, le mycélium dense montre des hyphes réguliéres, x 1,8-3-(4)
jm, 4 paroi mince, sans boucles et de nombreuses gloeocystides au con-
tenu dense ou pailleté. Plus profondément on observe encore des
éléments gloeocystidiens, mais moins nombreux et de forme plutdt nodu-
leuse.
Boucles: absentes
Cytologie: les cultures polyspermes sont constituées d'hyphes aux
articles réguliérement binucléés
Oxydases: acideeallique: +44++,0 ~galacol<* +4750
pi cresol 2 7 — tyrosine: +++,0
CODE: 6 = 7 =) 15% = 35) =86 = 938 -..47 = 53) o4 Oo
Remarque. Cette espéce qui exhale une nette et constante odeur agréa-
ble est caractérisée par sa croissance extrémement lente, ses nombreu-
ses sulfocystides irréguliéres sans schizopapilles, la présence
d'arthrospores (surtout visibles en culture sur lame) et son comporte-
ment hétérothalle sans boucle.
Si les mycéliums des Epithele et Pteridomyces ont une croissance
aisée, les germinations des spores en semis clairs ont été beaucoup
plus difficilesa obtenir avec des spores qui certes avaient voyagé
plusieurs jours. Notons toutefois que E. nikau et guadelupensis sont
homothalles lents, E. efibulata, parthénogénétique et P. lacteus,
hétérothalle. La préserice de laccase laisse supposer que toutes les
espéces sont a pourriture blanche quant 4 Vesiculomyces epitheloides,
il est hétéroth alle sans boucles, comme le type du genre V. citrinus,
ainsi que V. lactescens et ses gloeocystides sulfoaldhéydes trés
positives en culture contiennent du stearyl-velutinal (K. Gluchoff-
Fiasson, viva voce).
REMERCIEMENTS
Nos remerciements s'adressent tout d'abord 4 notre ami Gérard Gilles
a qui nous devons de nombreuses récoltes africaines et malgaches citées
icl, accompagnées de sporées, de notes sur le frais, avec données
microscopiques. Nous exprimons, en outre, notre gratitude envers les
directeurs du Jardin Botanique de 1'Etat de Bruxelles et de 1'Herbier
de Kew, nos collégues R.A. Maas Geesteranus et A.L. Welden pour leur
lecture critique de notre manuscrit , et 4 Monsieur H. Romagnesi pour
les nombreuses diagnoses latines incluses ici et, pour lesquelles il
a bien voulu nous faire profiter, une fois encore, de sa maitrise de
cette langue.
497
SUMMARY
We have studied the effused and epitheloid fungi collected in warm
countries. The fourteen species described in this paper cannot be
included in the four genera Epithele (Pat.) Pat. (1900). Epithelopsis
Julich (1976), Pteridomyces Julich and Skeletohydnum Jiilich (1979), as
definited by Julich. reais 6 ia olin
As already noticed by Patouillard (1899), sterile pegs occur in some
tremellaceous Heterobasidiomycetes (genus Heterochaete), and also in
a few Homobasidiomycetes such as Mycobonia, Veluticeps and Epithele
(sensu lato). We shall write pals eaboael ene last named genus, i.e.
effused species with indeterminate margins. To gather all these
effused species described nowadays in one genus would make this genus
dissimilar, heterogeneous and unworthy. We propose to place in the
genus Vesiculomyces Hagstrom (1977) emend., segregated from the genus
Gloeocys tidiellum Donk, the Malagasy Vesiculomyces epitheloides with
amyloid spores and sulfoaldehyde positive gloeocystidia. We synonymise
Megalocystidium (type M. leucoxanthum (Bres.) JUlich 1978) with
Vesiculomyces (type V. citrinus (Pers.) Hagstrom), because these two
genera contain some species of Gloeocystidiellum with smooth spores,
with or without clamps, with gloeocystidia habitually exhibiting a
positive sulfoaldehyde reaction (the gloeocystidia of V. citrinus
react strongly in cultures, but appear empty in fructification) .
As a consequence some combinations are proposed.
The species with non-amyloid spores do not form a homogeneous group.
Julich (1976 and 1979) created successively several monospecific
genera: Epithelopsis, Pteridomyces and Skeletohydnum. Their defini-
tions are based on the cyanophily of the spores, their mono- or dimi-
tism, and the size of the basidia. We have not collected species with
cyanophilous spores (genus Epithelopsis), known only from New Zealand
and Zaire (Ryvarden 1978), but the size of the basidia, as well as
the origin and development of a second hyphal system are progressive
characters, and we wondered if the study of new species would not
permit us to discover all kinds of intermediates. Our collections can
be separated easily in two groups according to the size of basidia
and the size of the spores: one group has narrow club-shaped basidia
(narrower than 5 \m) and spores shorter than 8 ym; the other group has
larger basidia, not club-shaped, and spores longer than 10 wif. But in
these two groups, there are species without skeletal hyphae (monomitic)
and species with skeletal hyphae more or less differentiatedand more or
less numerous. Table I shows the different localizations possible of
the skeletal hyphae, -when they exist-, in the group with large spores:
1) only in pegs (should we call these species dimitic?), 2) in the pegs
and in the context, 3) in the context only.
In the light of these facts it is necessary to either create several
new genera with an absolute care for homogeneity (as it appears some
people with probably a desire to leave their names to prosterity have
done!) or accept, until a more complete knowledge of the epitheloid
fungi of the world, that the kinds connected by the means of interme-
diates should be gathered again into a small number of less homogenous
498
genera. We agree with the second saqlution, and we includespecies
with skeletal hyphae in the genus Ptéridomyces emend., and we
maintain all the large spored species, whatever their mitism should be,
in the genus Epithele. We place also in the genus Epithéle an unclamped
species (E. efibulata sp. nov.). Three new African species of Pterido-
myces, which are not growing on ferns, are proposed (P. P. lacteus, rose-
olus, sphaericosporus), and six new Epithele species (E. citris) citrispora, |
efibulata, uadelupensis, macarangae, malaiensis, Seal i eporae Four
of these are African and most of them grow on palm-trees. Lastly, we
described Vesiculomyces epitheloides from Madagascar.
We give cultural characters of seven species: E. efibulata is parthe-
nogenetic without clamps. Such a species must not be placed ina
different genus from the one with clamped species. E. nikau and
guadelupensis are homothallic and clamped. Vesiculomyces epitheloides
ls heterothallic without clamps as are V. citrinus and V. lactescens.
BIBLIOGRAPHIE
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BOIDIN J., LANQUETIN P., TERRA P. et GOMEZ C.E. - 1976 - Vararia subg.
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CORNER E.J.H. - 1953 - The construction of Polypores. Phytomorphology
Se elo2= 167 »
DONK M.A. - 1957 - The generic name proposed for Hymenomycetes VII.
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HAGSTROM E. — 1977 — Vesiculomyces Hagstrom gen. nov. segregated
from Gloeocystidiellum (Corticiaceae) Bot. Noe. 1130: p53 542
HALLENBERG N. —- 1980 - New taxa of Corticiaceae from 1} N. Iran (Basidio-
mycétes). Mycotaxon 11: 447-475.
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mycétes) XI. Boidinia, a new genus segregated from Gloeocystidiellun.
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Corticieen II Sitz. Kaiserl. Akad. Wiss. Math. Nat. Kl. 116: 739-
782.
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genera and species. Persoonia 10: 137-140;
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MYCOTAXON
VOLT ENV None pp, 6500-500 January-March 1983
NEW SOUTH AMERICAN RESUPINATE POLYPORES |
Mario Rovichenc ence
Departamento de Ciencias Bioldgicas
Facultad de Ciencias Exactas y Naturales
Universidad de Buenos Aires
Argentina
During a survey of the resupinate polypores) in Argentina,
two species pertaining to Amylosporus Ryv. and Junghuhntia
Corda and a new variety of Skeletoecutts nivea (Jungh.)Keller
were found which are here described as new.
METHODS
Microscopic examinations of basidiocarps were made from
freehand sections mounted 1n 5% KOH aqueous phloxine i
Melzer's reagent (Singer, 1949:80) and in 0,5% cotton blue
in 25% lactophenol. Drawings were made with a camera lucida.
Herbarium abbreviations are from Holmgren § Keuken (1974).
DESCRIPILON OP oY BCLES
AMY LOSPORUS. WRIGHTLT sp: i nov. (Fi geass)
Bastdtocarpus annuus vel perennis, late resuptnatus, roseo-ltlact-
nus, gossyptnus vel suberosus, non xanthochrous, contextus vinaceus.
Systema hypharum dimtttecum. Hyphts generattvts duorum typt: alaftbulatts
tenutbus tnerassatts, 2,1-10,5 pm dtam.; b)septatus unt vel plurtfrbula-
tis vertictllatisque, 5,2-10,4 pm dtamn., ramosts aftbulatis. Hyphis ske-
Lettbus, 2,6-5,7 ym dtam., partetts tnerassatts hyalints vel castanets,
dextrtnotdets ecyanophtltsque. Hymento bastdits aftbulatts septitts basa-
lts, clavtformtbus et brevt sttpttatts, tetrasports. Bastdtosports late
elltpsotdets vel subglobosts, aptculatts, tenut asperulatis, 3,1-3,6 x
2,6 pm, amylotdes, acyanophtlts. Cysttdtolts fustformtbus adsunt. Pu-
trefacttone ltgno tgnota.
Holotypus: Argenttna, Mistones, Parque Nactonal Cataratas del Igua-
zu, versus viam ad Apept, leg. M.Rajchenberg M-3503, 4/III/1982, tn her-
barto BAFC n© 28210 conservatus est.
A mycologo argenttnensts J.E. Wright haee spectes dicata est.
1.This paper is part of a project on xylophilous Basidiomycetes subsidi-
zed by a grant from the Consejo Nacional de Investigaciones Cientifi-
cas y Técnicas, Argentina. It is part of a thesis submitted to the
University of Buenos Aires as partial fulfillment of the requirements
for the degree of Doctor in Biology.
2.Fellow of the above Consejo.
501
FRUITBODY annual to perennial, widely effused, easily
separable from substratumand the annual layers from each
Other, 25 x 10 x 0,5 cm, cottony to corky. Margin wide,
irregular, velvety. Pore surface lilaceous pink when fresh,
turning cream or yellowish cream when dried. Pores 7-8 per
mm. Context lilaceous when fresh, turning dark chestnut or
vinaceous when dried; each layer up to 2 mm wide. Tubes
short, up to 1 mm long. Alkali reaction negative with 5% KOH.
HYPHAL SYSTEM dimitic. Generative hyphae of two types:
a) clampless, branched and thin walled hyphae, 2,1-10,4 pm
wide;e.ound in all ‘the truitbody (Fig. 2); and b) with one
OGemubciple "clamps; branched or not, 5,2-10,4 pm wide, with
Cclampless and narrower branches, only found in the context
(Fig. 3). Skeletal hyphae abundant, scantily to profusely
branched, 2,6-5,7 pm wide, with thickened hyaline or chest-
nut walls, cyanophilous and variably dextrinoid in Melzer's
reagent (Fig. 4). BASIDIA claviform to shortly pedicellate,
ieee X 5,/-0,8 pm; tetraspored, not clamped at base
(Fig. 5). BASIDIOSPORES broadly ellipsoid to subglobose,
thin to slightly thick walled, minutely asperulated (when
observed in Melzer's reagent, Fig. 5, and SEM, ao ee rb ie
auyrora, noncyanophilous:, 3,1-3,6.x 2,6 pms -Fusitorm cysti-
dtolsspresent; cystidia’ absent.
HABITAT: on unidentified angiosperms and associated with
an unknown rot. DISTRIBUTION: in the subtropical rain forest
of Misiones province.
Loe.)
SEM micrographs of
basidiospores of
Amylosporus wrtghttt
MATERIAL STUDIED: Argentina, Misiones, Colonia General Belgrano,
Estacion Forestal, leg. Wright, Deschamps and del Busto, 29/X/73 (BAFC
28211); Parque Nacional Cataratas del Iguazi, leg. M.Rajchenberg M-3503,
4/111/82 (BAFC 28210, holotype).
REMARKS: In spite of macromorphological differences with
the type species of the genus Amylosporus Ryv., Tyromyces
gramtntecola Murr. (=Amylosporus campbellt (Berk.)Ryv., fide
Ryvarden, 1977:217) which is pileate and centrally or latera-
ily stipitate, the peculiar hyphal system with two types of
septation on the generative hyphae, the asperulated amyloid
Fig. 2-5, Amylosporus wrtghtit: 2.simple septate generative hyphae;
3.simple and multiple clamped generative hyphae, unbranched or with sim-
ple septate branches; 4.skeletal hyphae; 5.hymenium and basidiospores.
Fig. 6-10, Junghuhnta microspora: 6.generative hyphae; 7.skeletal
hyphae; 8.hymenium, cystidia and basidiospores; 9.cystidia (without
crystals); 10.pore mouth with incrusted hyphae and cystidia.
504
Spores ands tue peciceblaterGlLamphess basidia> PON t-4OuUt eno
the) congenerity of this mew Speciés with ‘the latter eine
skeletal hyphae with cyanophilous and variably dextrinoid
walls relate this species to Wrightoporta Pouz. which dif-
fers in having generative hyphae and basidia with only sim-
ple clamped-septa:
JUNGHUHNIA MICROSPORA sp. nov. (Fig. 6-10)
Bastdtocarpus annuus, resuptnatus, suberosus vel ftbrillosus, cre-
meus, late castaneus vel fuscus, non xanthochrous; ports 5-7-(8) per mn.
Systema hypharum dimttteum. Hyphts generativis ftbulatts, 2,1-5,7 pm
diam. Hyphts skelettbus 2,1-6,8 ym dtam., partettbus tnerassatts lutets.
Hymento cum bastdits clavtformtbus, tetrasports. Bastdtosports elltpsot-
deis, hyalinis, dqpteulatts, parietibus angustis olaprts, (2,0-3,00enee
1,6 pm, tnanylotdes, acyanophilts. Cysttdits tnerustatts adsunt, copto-
sts extremttattbus diseptmenttts. Putrefaettone ltgno alba.
Holotypus: Argentina, Mistones, Parque Nacional Cataratas del Igua-
Zu, versus viam ad Apepu, leg. M.Rajchenberg 4/III/1982, tn herbarto
BAFC n@ 27993 conservatus est.
FRUITBODY annual, resupinate, totally attached ito scups—
trate, corky to fibrillose. Margin present or not ~myec mc
or velvety. Pore surface cream’ colour With light) che2ria.
tints when fresh, turning. beige, orange-coloured)orscneseaut
upon drying, and with the pore mouths remaining ash-coloured.
Pores -5-7-(8) perimm, round’ to, Slightly angular. Contes:
thin, cream coloured, up to 0,5 mm wide. Tubes up to 3 mm
long. Alkali reaction negative with 5% KOH.
HYPHAL SYSTEM -dimitic>’ Generative hyphaeclamped,
hranched, hyaline, walls thin or slightly thickened, 2,1- 9.7
pm wide (Fig. 6). Skeletal hyphae unbranched, thick-walled;
2, b70,8,pm wide (Pig. 7). HYMENIUM 9,4-1354 um thicks (hone
BASIDIA. claviform, -8,5-10,4 x 4, lh4,7 um. tet raspoueds
BASIDIOSPORES ‘ellipsoid, hyaline; thin-walled, 2,0-3;,6 oot.
1,6 pm, inamyloid, acyanophilous. CYSTIDIA few to abundant,
clavate. (Fig. 9), 4,2-10,4 pm wide,» thick-walled, vembeddea
in the trama or projecting up to-.30 pm, abundant in theppore
mouths (Fig. 10), heavily incrusted apically, arising from
skeletal hyphae.
HABITAT: on angiosperms, associated with a white foc
DISTRIBUTION: tn the, subtropical rain forest of Misrones
province.
MATERIAL STUDIED: Argentina, Misiones, Parque Nacional Cataratas
del Iguazi, leg. M.Rajchenberg 5/III/80 (BAFC 27992); ibid, leg. ipse
4/T11/82, M-3504 (BAFC 27993, holotype), M-3510 (BAFC 27994) and M-3511
(BAFC 27995).
REMARKS: Junghuhnta mtcrospora differs from other species
of the ‘genus by its small pores. and spores. J: ‘collabene
(Fr.)Ryv. has a typical brick-red pore surface and larger
3.Tyromyces graminicola Murr. (holotype studied, NY) also lacks clamps
at the base of the basidia in the fruitbodies. Gilbertson & Lowe
(1970:704) in their study of this species do not give information on
this feature, but illustrate simple clamped basidia produced in culture.
505
Wimaneoral to pcylindrrc“sporesmimye x 1.27, pM) a. netida
(Pers::fr.jJRyv. differs in having Jarger ellipsoid spores
(3,5-4,5 x 2,0-3,0 pm). Porta novae-zelandtae G.H. Cunn has
Simitar pore and spore sizes, but skeletal hyphae are varia-
Diywiperusted and never form Cruescystidia; besides, a siiumy
Supstance embeds all hyphal elements which is lacking in
Junghuhnta mtcrospora.
SKELETOCUTIS NIVEA (Jungh.)Keller var. DILUTA var. nov.
A typo dtffert hyphits skelettbus dtffluenttbus tn 5% KOH et levt-
ter amylotdets.
Holotypus: Argentina, Mtstones, Iguazu, Puerto Esperanza, in satto
Alto Paranda, leg. M.Rajchenberg III/1982, tn herbarto BAFC n@ 28250
conservatus est.
FRUITBODIES and microscopical features are similar to
tise. on the typical variety . 1, SKELETAL HY PRAGY da Drermyrada-
Cally begs SssOlviange an .5% \KOH: “and\in ‘exhibiting a weakly
amyloid reaction of their walls.
BAcinagl: Only found On Pinus taeda Jogs and other
unknown gymnosperms, associated with a white rot. DISTRIBU-
TION: only found in subtropical implanted woods in Misiones
DLrovances
MATERIAL STUDIED: Argentina, Misiones, IguazOi, Puerto Esperanza,
Alto Parana plantation, leg. M.Rajchenberg III/1982 (BAFC 28250, holo-
type); ibid., leg. S.Blumenfeld 25/III/81 (BAFC 27657).
REMARKS: The remarkable character of this variety has
only been known up to now in Porta ctnerascens (Bres. ex
MEracs. Noacc., a Species that does not haye an adequate ge-
Neticmatspostt1on. Ryvarden (1976:°82) placed’ at in’ Antrodia
Mero eewras 4. it naplcart (Berk: 6 Br. ex Berk.)Ryv. (basio-
nyNeroevyporus lindbladtt Berk. G BY. .ex Berk.) but the type
Of wood-rot. and cultural characters (Nobles, 1965:1132; Na-
kasone, 1981:108; and own unpublished data) show notable
differences with those of Antrodta species (when compared
with the table published by Domafiski, 1969:67). Whether the
Gapacaty, ofl dissolving the walls in KOH solution may be ‘a
Cilcm@eOurelate Portia cineraseene With Skeletocutia nivea
Veter rura is Still a pending question; since P. eznerge=
eens lacks typical incrustations present in Skeletoecutis
(=“Inerustoporia Dom.) species (as shown by Keller, 1979:347).
Besides, cultural features are different, as shown by a com-
Parasom Of the principal code numbers, (fide Nobles scheme,
ioGeeioue 2a. 7, 1n,Poria eineraseens, ,and) 2. 3.) (8). Oo.
Minoweretocutite species (David, 1971795; Nobles, 196527129
Bnomota pers, »h978:95). Notwithstanding this, there, is) asre-
markable chemical correlation between presence of dissolving
walls and amyloidity.
4.This striking reaction may be well observed by mounting the sections
in phloxine solution and adding one drop of KOH solution while looking
through the microscope. Walls evanesce, actually dissolve, and only a
cellophane-like outer pellicle remains. The walls do not reconstitute
when KOH solution is replaced overnight with phloxine solution.
ACKNOWLEDGMENTS
I wish to express my gratitude to Dr. Ma. Larsen. (mae
dison) and Drs. LoaRyvarden (Oslo), for ‘critreally sreading
the M.S. and making invaluable suggestions. Particularly
gratifying were the discussions held with the latter.
REFERENCES
DAVID, A., 1971. Caracteres myceliens d'Incrustoporta percandida Mal. &
Bert. Acta’ Phytotax.sBarcinon. 8:95-97.
DOMANSKI, S., 1969. Wood-inhabiting fungi of Bialowieza virgin forests
in Poland. VI. Antrodta ramentacea (Berk. & Br.)Donk. Acta. Soc.
BOt. Poloniges58.57-08.
GILBERTSON, R.L. & LOWE, J.L., 1970. Tyromyces gramintcola in North
America. Mycologia 62:699-706.
HOLMGREN, K.P. §& KEUKEN, W., 1974. Index Herbariorum. Regnum Vegetabile
O25-597 pp:
KELLER, J., 1979. Ultrastructure des hyphes incrustées dans le genre
Skeletocutts. Persoonia 10:347-355.
NAKASONE, K.K., 1981. Cultural studies on Porta cinerascens, P. rivulo-
sa and P. subvermtspora (Aphyllophorales, Basidiomycotina). Myco-
taxon 1 oe105—00 1:
NOBLES, M.K., 1965. Identification of cultures of wood-inhabiting Hyme-
nomycetes., Can. J. Bot. 45% 1097-11139.
RYVARDEN, L., 1976. The Polyporaceae of North Europe, Vol. I. Albatre-
llus-Inerustoporta. Fungiflora, Oslo, 214 pp.
--- 1977. Type studies in; the Polyporaceae_10. Species described by
J.M. Berkeley, either alone or with other authors, from 1844 to
IBS 2NOTWs ds. BOte 2422) o=Z00.
SINGER, R., 1949. The Agaricales in modern taxonomy. Lilloa 22, 832 pp.
STALPERS, J.A., 1978. Identification of wood-inhabiting Aphyllophorales
in pure culture. Studies in Mycology n°16, 248 pp.
MYCOTAXON
VOUSRAV I, Node2, pp. 507-506 January-March 1983
COLLETOTRICHUM GLOEOSPORIOIDES (PENZIG) PENZIG ET SACCARDO
S. R. PENNYCOOK
Plant Diseases Division, DSIR, Auckland, New Zealand
The authorities for the anamorph of the common plant
pathogen Glomerella cingulata (Stoneman) Spaulding et
Schrenk have been variously, but incorrectly, cited. The
correct nomenclature is:
Colletotrichum gloeosporioides (Penzig) Penzig et Saccardo,
in Penzig, Note micologiche. III. Seconda contribuzione
allo studio dei funghi agrumicoli. Atti del Reale
Istituto Veneto di Scienze, Lettere ed Arti, Serie 6,
elo 6702 <b884-
Basionym: Vermicularia gloeosporioides Penzig, Funghi
agrumicoli. Contribuzione allo studio dei funghi
parassiti degli agrumi. Michelia 2: 450. 1882.
Penzig employed the combination 9 times in his 1884 paper,
but only 2 of these occurrences refer to a previously
published description as required for valid publication of a
new combination (International Code of Botanical Nomen-
Crature;e 1975, Article) 32) 2. The first of these, on p. 670
under "Colletotrichum gloeosporioides Penz. et Sacc.", not
only indicates the basionym but also includes a taxonomic
discussion and a redescription of the species.
Penzig's 1882 paper was also distributed as a separatum
with its own pagination (see Lindau & Sydow, 1909, item
20412), so that the basionym has been cited as published in
EPungeeagre;, p. 66. (e.g. by Penzig, 18847 7p. 671). Penzigs
1884 paper may also have been distributed as a separatum;
tirisewould account. for Saccardo'’s (1884, p.. 735); citation of
the new combination as published in "F. Agrum. II. p. 6."
Arx (1957a, p. 427) also cites the publication in this way,
but assigns incorrect dates to both the basionym and the new
combination. [Arx's annotation "teste Shear et Wood, 1913"
508
does not assist, since Shear & Wood (1913, pp. 27, 104)
refer only to Penzig's 1882 paper.] These errors have been
perpetuated by Arx (1957b, p.' 38; 1970, p.) 159): and Sutton
(1980, p. 530). The correct dates of Penzig's two papers are
defined by the journal title pages and the author's prefaces
("1 Decembr. 1882" - "Giugno 1882"; and "dal Novembre 1883
all' Ottobre 1884" - "Gennajo 1884", respectively).
REFERENCES
Arx, J.A. von (1957a) Die Arten der Gattung Colletotrichum
Cda. Phytopathologische Zeitschrift 29: 413-468.
Arx, J.A. von (1957b) Revision der zu Gloeosporium
gestellten Pilze. Verhandelingen der Koninklijke
Nederlandse Akademie van Wetenschappen, afd.
Natuurkunde, 2de reeks, 51(3): 1-153.
Arx, J.A. von (1970) A revision of the fungi classified
as Gloeosporium. Bibliotheca Mycologica 24: 1-203.
Lindau, G.; Sydow, P. (1909) Thesaurus litteraturae
mycologicae et lichenologicae, vol. 2. Leipzig.
Penzig, O. (1882) Funghi agrumicoli. Contribuzione allo
studio dei funghi parassiti degli agrumi.
Michelia 2: 385-527.
Penzig, O. (1884) Note micologiche. III. Seconda contri-
buzione allo studio dei funghi agrumicoli. Atti del
Reale Istituto Veneto di Scienze, Lettere ed Arti,
Serie 6, 2(5): 665-692.
Saccardo, P.A. (1884) Sylloge fungorum omnium hucusque
cognitorun; “vol. Ili.) Padova:
Shear, C.L.; Wood, A.K. (1913) Studies of fungous parasites
belonging to the genus Glomerella. US Department of
Agriculture, Bureau of Plant Industry, Bulletin 252:
T1220:
Sutton, B.C. (1980) The coelomycetes. Commonwealth
Mycological Institute, Kew.
MYCOTAXON
Viltewmen Vtts NO.) 25 DD. SUOso Lo January-March 1983
ECTOMYCORRHIZAE OF SELECTED CONIFERS
GROWING IN SITES WHICH
SUPPORT DENSE GROWTH OF BRACKEN FERN
JAN ACSAI
Department of Plant Pathology,
Colorado State University, Fort Collins, Colorado 80523
and
DAVID L. LARGENT
Department of Biological Sciences,
Humboldt State University, Arcata, California 95521
SUMMARY
Fifteen ectomycorrhizae are described for Abies concolor of which the mycobiont is
known for six and thirteen ectomycorrhizae are described for Pseudotsuga menziesii, of
which three mycobionts were identified. The presence of these mycorrhizae on conifer
seedlings located in sites overgrown by bracken fern suggests that the phytotoxins of
bracken fern have little effect on ectomycorrhizae formation in nature in these areas.
INTRODUCTION
Many acres of formerly forested land in the Pacific Northwest now support dense
growth of bracken fern, Pteridiuwm aquilinum (L.) Kuhn (Isaac, 1940). One of the reasons
bracken fern can dominate vegetation is its ability to produce allelopathic chemicals
(Glass, 1976; Gliessman, 1976; Gliessman and Muller, 1972).
Bracken fern produces four phytotoxic phenolic acids: p-hydroxybenzoic acid, vanillic
acid, ferulic acid, and p-hydroxycinnamic acid (Glass, 1976; Glass and Bohm, 1969). Small
amounts of the toxins can be found in the rhizome but the highest concentrations are found
in the fronds (Gliessman and Muller, 1972). Each fall, the current year’s growth of fronds
die and the toxins are released by rainfall into the soil as free phenolic acids (Glass, 1976;
Whitehead, 1964). Whitehead (ibid) found that in sandy soil under bracken fern the four
phenolic acids were present in the following concentrations: p-hydroxybenzoic acid, 3.9 x
10-5 M; vanillic acid, 4.9 x 10> M; ferulic acid, 0.4 x 10-5 M; and p-hydroxycinnamic acid,
4.2 x 10° M. Once in the soil they can be absorbed from the soil solution by plant roots
(Glass, 1976; Gliessman and Muller, 1972).
Gliessman and Muller (1972) discovered that radicle growth of Bromus rigida Roth. is
inhibited by water extracts of the fern fronds. They also found that shoot growth of Avena
fatua L. is inhibited when the plants are grown in soil containing crumbled fronds. In
another experiment, the four phenolic acids, in concentrations similar to those found in the
soil by Whitehead (1964), were added to hydroponically grown barley, Hordeum vulgare
L.C.V. Carlsberg & C.V. Zephyr, resulting in an overall decrease in root volume (Glass,
1976).
50
Ectomycorrhizae benefit their plant associates in many ways. The fungal partner of
the association can increase the nutritional uptake of the plant by expanding the volume of
soil the plant can exploit (Bowen, 1973; Harley, 1959). Another benefit of the mycorrhizal
association for the tree is protection of the root against toxic chemicals. Mycorrhizae are
considered essential in the revegetation of soil containing residual heavy metals from
anthracite mining (Schramm, 1966). Toxins in bituminous mining spoils either did not
affect, or increased, the growth of trees with mycorrhizae (Medve et al., 1977). The herbi-
cides, 2,4-D, 2,4,5,-T, and amitrole, and the pesticide malathion do not affect growth of
mycorrhizal fungi in culture (Dasilva et al., 1977). Simazine, a weed control herbicide,
does not affect mycorrhizal development on Pinus seedlings (Smith and Ferry, 1979).
Ectomycorrhizae also protect their associates against effects of air pollution. Carney et al.
(1978) demonstrated that loblolly pine (Pinue taeda L.) with ectomycorrhizae were more
resistant to the effects of sulfur dioxide and ozone than those seedlings without
mycorrhizae.
Because mycorrhizae are known to increase the volume of soil exploited by a plant and
because they are able to tolerate certain toxic effects, some mycorrhizae may enable
conifer seedlings to survive among bracken fern. This project describes an attempt to
explore the hypothesis that fungi form ectomycorrhizae with conifer seedlings on sites on
which bracken fern are growing, and that these ectomycorrhizae protect seedlings against
effects of the fern phytotoxins.
MATERIALS AND METHODS
Five sites in Humboldt County, California, were selected on the basis of presence of
bracken fern and conifer seedlings and differences in elevation, slope aspect, parent mate-
rial, and conifer species (Table I). A comparison of the features of the five sites shows a
variation in elevation from 880 to 1370 meters, and a great diversity in slope aspects.
There were three parent material types on the sites. Two were igneous in origin; the
peridiotite-gabbro complex of sites I and II and the diorite-monorite-granite combination
of site V. On sites I and II there is an additional component, serpentine outcrops. The third
parent type, sandstone and shale on sites III and IV are sedimentary in origin. The vegeta-
tion of all five sites included Pseudotsuga menziesii (Mirb.) Franco. but differ in associated
tree species. Abies concolor (Gord. & Glend.) Lindl. is an additional component on sites I, II
and IV, Pinus ponderosa Laws. is found on sites II and IV, and Libocedrus deurrens is on
sites I and II.
Mycorrhizal roots were collected from May to December 1979, and subsequently
described in the laboratory. Descriptions were based on characters of the mantle, Hartig
net, attached rhizomorphs, attached mycelium, and the rootlet. Mycorrhizae that were
very similar in a number of characters were combined. Some of the fungal partners are
taxonomically unknown because there were no fruiting bodies found in contact with those
roots. Although no sclerotia of Cenococcum graniforme (Sow.) Ferd. & Winge were found,
it was identified by its characteristic appearance of jet black mycelium. Voucher speci-
mens of the roots are located in the Humboldt State University Cryptogamic Herbarium
and stored in vials of 50:50 (v:v) mixture of Craf 3 solution (9:1, v:v, chromium trioxide:
glacial acetic acid) and 20% formalin.
In both the spring and fall of 1979, fruiting bodies of fungal species were collected
from the sites. Three criteria were used to determine mycorrhizal associations. One, as
each fruiting body was found it was examined to determine if there was physical contact
with or a close proximity to mycorrhizal roots. Two, macroscopic and microscopic features
of the hyphae at the sporocarp base were compared to features of the hyphae attached to
the mycorrhiza. Three, the literature was reviewed for reports of a mycorrhizal associa-
tion between the fungal species and the conifer species. The sporocarps were dried and
stored as voucher specimens.
Sala
jUasqe SuUOI}O9UUO0D duIBID = 0-
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On each site one or two root samples consisting of sixty to one hundred rootlet tips
from each of twenty randomly selected seedlings were collected. Samples were collected
from seedlings of Abies concolor, Pseudotsuga menziesii, and Pinus ponderosa. Each tip
was examined and the percentage of all tips that were mycorrhizal was calculated. Tips
were considered to be mycorrhizal if they were swollen, had no root hairs, were shortened,
and had visible signs of hyphae on the rootlet surface. Questionable mycorrhizal roots were
dissected and examined under a microscope.
RESULTS AND DISCUSSION
Examination of root samples from the field sites showed that over 90% of the roots had
mycorrhizal infections on all sites. The average infection of the root samples from each site
was as follows: site I, 99.7%; site II, 99.1%; site III, 98.1%; site IV, 99.4%; and site V, 91.4%.
All mycorrhizae described were ectomycorrhizae associated with either Abies concolor
or Pseudotsuga menziesu.. Except for Cenococcum graniforme, the fungal associates of
Abies concolor mycorrhizae displayed different characteristics from those of Pseudotsuga
menziesii mycorrhizae. A variety of mycorrhizal types found on the field sites exhibited
differences in rhizomorphs, mycelium attached to the mantle, the mantle, the Hartig net
and the rootlets. When compared, rhizomorphs differed in form, either ropy or ropy-wefty,
in color, from white to yellow, and in diameter. In all specimens examined, the mycelium
attached to the mantle consisted of hyaline hyphae that had either rough or smooth walls
and was 1.5 to 4 um in diameter. Characteristics of the mantle varied greatly in color and
texture. The mantle of all but two of the types examined consisted of parenchymatous
hyphae. Of the mantles made of synenchymatous hyphae, the mycobiont of one was Ceno-
coccum graniforme, the other was unknown. The Hartig nets analyzed were composed of
one to several hyphae between the cells of the root cortex and extended to the endodermis
or to one cell layer from the endodermis. Rootlets examined were shortened and swollen
but differed from one another in branching patterns, from monopodal to pinnate, remi-
form or tuberculate (Tables II & III).
The variety of mycorrhizal forms and mycorrhizal infection of over ninety percent on
all sites indicate that formation of mycorrhizae by some fungi on the rootlets of Abies
concolor and Pseudotsuga menziesii is not inhibited by the phytotoxins produced by
bracken fern.
ACKNOWLEDGMENT
This work was supported by Cooperative Grant #54-PNW-78 with the Pacific Northwest
Range and Experiment Station, Portland, Oregon.
LITERATURE CITED
Bowen, G.D. 1973. Mineral nutrition of ectomycorhizae. Pp 151-205. In: Ectomycorrhizae,
their ecology and physiology. Eds., G.C. Marks and T.T. Kozkowski. Academic Press,
New York.
Carney, J.L., H.E. Garrett, and H.G. Hendrick. 1978. Influence of air pollutant gases on
oxygen uptake of pine roots with selected ectomycorrhizae. Phytopath. 68: 1160-63.
Dasilva, E.J., L.E. Hendriksson, and M. Udris. 1977. Growth responses of mycorrhizal
Boletus and Rhizopogon species to pesticides. Trans. Brit. Mycol. Soc. 68: 434-387.
Glass, A.D.M. 1976. The allelopathic potential of phenolic acids associated with the rhizo-
sphere of Pteridium aquilinum. Can. J. Bot. 54: 2440-44.
Glass, A.D.M. and B.A. Bohm. 1969. The accumulation of cinnamic and benzoic acid
derivatives in Pteridium aquilinum and Athyrium felix-femina. Phytochem. 8: 371-
aT:
Gliessman, S.R. 1976. Allelopathy in a broad spectrum of environments as illustrated by
bracken. Bot. J. Linn. Soc. 73: 95-104.
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Gliessman, S.R. and C.H. Muller. 1972. The phytotoxid potential of bracken, Pteridium
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MYCOTAXON
VOR eNO. 12. DD 519-556 January-March 1983
MYCORRHIZAE OF ARBUTUS MENZIESIT PURSH.
AND ARCTOSTAPHYLOS MANZANITA PARRY
IN NORTHERN CALIFORNIA
JAN ACSAI
Depart».ent of Plant Pathology
Colorado State University, Fort Collins, Colorado 80523
and
DAVID L. LARGENT
Department of Biological Sciences,
Humboldt State University, Arcata, California 95521
SUMMARY
51 arbutoid-, 2 arbutoid-ericoid- and 7 ecto-mycorrhizae are described as occurring on
roots of either Arbutus menziesii Pursh. or Arctostaphylos manzanita Parry in 8 different
sites in northern California. All sites consisted of a predominance of Arctostaphylos man-
zanita with scattered trees of two conifers, Pinus ponderosa Laws. and Pseudotsuga men-
ziesti (Mirb.) Franco.
INTRODUCTION
Arbutus menziesii Pursh. and Arctostaphylos species growing in the shrublands and
woodlands of Northern California commonly form mycorrhizae (Largent et al., 1980b).
However, little research attention has been focused on the description of thee mycorrhizae
or identification of the fungal symbiont. Several pure culture synthesis studies have estab-
lished that various fungi which are known to form ectomycorrhizae with conifers can also
form arbutoid mycorrhizae with Arbutus menziesii, Arctostaphylos uva-ursi (L.) Spring.,
and Arctostaphylos manzanita Parry (Comes, 1980; Molina and Trappe, 1982; Zak, 1976a,
1976b). While there is no certainty that these mycorrhizae will form under natural condi-
tions, it demonstrates the variety of fungal species which are potential mycorrhizal
formers with these plants. By tracing the rhizomorphs and mycelia between sporocarps
and mycorrhizae on root samples collected in field work, three fungal symbionts have been
identified: Cortinarius aureifolius Peck var. hesperius J.-F. Ammirati in association with
Arctostaphylos uva-ursi (Zak, 1973); Cortinarius zakii J.F. Ammirati with Arbutus men-
ziesii (Zak, 1974); and Amanita gemmata (Fr.) Gill. with Arctostaphylos manzamta (Lar-
gent, et al., 1980a). A fourth fungus, Cenococcum graniforme (Sow.) Ferd. & Winge,
identified by the appearance of the mycorrhizae, associates with Arbutus menziesii and
several Arctostaphylos spp. (Largent et al., 1980b; Trappe, 1964; Zak, 1973, 1974).
The mycorrhizae formed with Arctostaphylos spp. and Arbutus spp. are pre-
dominantly arbutoid, but ericoid forms and ectomycorrhizae have also been observed
(Largent et al., 1980b; Mejstrik and Hadac, 1975; Trappe, 1964; Zak, 1973, 1974). The type
of mycorrhizae formed by Cenococcum graniforme has been placed in more than one cate-
gory. Field studies show that this fungus is capable of forming ectomycorrhizae with Arc-
520
tostaphylos and Arbutus (Largent et al., 1980b, Mejstrik and Hadac, 1975), while in pure
culture experiments it develops into an arbutoid type (Molina and Trappe, 1982; Zak,
1976a, 1976b).
Because little field work has been done on the mycorrhizae of Arctostaphylos spp. and
Arbutus spp. this study was undertaken to provide more complete information on the char-
acteristics of their mycorrhizae and to identify some of their natural fungal symbionts.
MATERIALS AND METHODS
Root specimens were collected from Arbutus menziesvi and Arctostaphylos manzanita
on three sites in Humboldt County, California, from 1979 to 1981. The predominant vege-
tation on the sites is Arctostaphylos manzanita, with scattered Arbutus menziesii, Pinus
ponderosa Laws., and Pseudotsuga menziesvi (Mirb.) Franco. One of the sites also has Quer-
cus garryana Dougl. and Quercus chryolepis Liebm.
The roots were examined both macroscopically and microscopically and described
using features of the mantle, attached mycelium, attached rhizomorphs, and the rootlets.
The root specimens examined are stored in Craf #3 solution at Humboldt State University,
Arcata, California.
Sporocarps of fungi associated with the mycorrhizae were also collected whenever
observed. Fungal species were determined to be mycorrhizal symbionts if the sporocarps
were in physical contact with or repeated close proximity to mycorrhizal roots, and the
features of the mycelium or rhizomorphs at the sporocarp base were identical to those of
the mycorrhizal root. The sporocarps were dried and are stored as voucher specimens in
the Humboldt State University fungal herbarium.
RESULTS
Mycorrhizae were found on all but one of the root samples collected. Repeatedly, a
number of different mycelial and rhizomorph types were noticed in the soil, concentrating
in the area surrounding a rootlet complex. Root samples often had up to five different
mycorrhizae on a 10-20 cm length of root.
Fifty-five mycorrhizae were described in association with Arctostaphylos manzanita
and five with Arbutus menziesii. Most of the mycorrhizae examined, including Cenococcum
graniforme, were arbutoid (Tables I-IV). Anatomically, these mycorrhizae were all very
similar, with well developed mantles, a Hartig net extending between cells of the outer-
most layer of cortical cells, and intracellular hyphae packed into the cells of the first layer
of the cortex. The morphology of the specimens was highly variable, differing in the struc-
ture, texture and color of the mantle, attached mycelium and rhizomorphs, and the
branching forms of the rootlet complexes. A second type of mycorrhiza examined was
intermediate between ericoid and arbutoid forms (Table V). These roots exhibited a thin
mantle with a weak or nonexistent Hartig net, intracellular hyphae in the cells of the first
one to two layers of cortical cells, and typical arbutoid swelling and branching patterns of
the rootlets. Roots determined to be ectomycorrhizal differed from the arbutoid types only
in the absence of intracellular hyphae (Table VI). The Hartig net, unlike those found in
ectomycorrhizal conifer roots, extended only between the outermost one or two layers of
cortical cells.
Several fungi were identified as mycorrhizal associates of Arctostaphylos manzanita.
Species of Cortinarius were most common: Cortinarius zaki, Cortinarius fibrillosipes
var. fibrillosipes J.F. Ammirati, and two unidentified species. A species of Calocybe and
Boletus aereus Bull. ex Fr. were also identified. Cenococcum graniforme, identified by the
characteristics of the mycorrhizae and the presence of black sclerotia, was found in associ-
ation with both Arbutus menziesii and Arctostaphylos manzanita. Lyophyllum sp. was the
only other fungus identified in an association with Arbutus menziesii.
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536
DISCUSSION
The amount of mycelial and rhizomorph development in the soil surrounding the roots
of Arctostaphylos manzanita and Arbutus menziesii and the abundance and variety of
mycorrhizae described, support the hypothesis put forward by Molina and Trappe (1980)
that mycorrhizal roots of ericaceous plants may act as a reservoir of fungi that can be used
by conifers in forests that are becoming re-established.
Harley (1959) proposed that arbutoid mycorrhizae are intermediate between
ectomycorrhizae and ericoid mycorrhizae. Molina and Trappe (1980) hypothesized that the
arbutoid form is a type of ectomycorrhiza. The results of the research done in this paper
substantiate the theory of a close relationship between different types of mycorrhizae. The
difference between types of mycorrhizae may be caused by any number of environmental
factors tht affect the ability of the fungus to physically penetrate the cells of the cortex or
affect the production of certain fungal or plant hormones. A gradient of environmental
factors could result in a number of intermediate forms between two recognized categories
of mycorrhizae. The arbutoid-ericoid forms described here and observed by Largent et al.
(1980b) illustrate these intergrading forms.
Further research consideration should be given to the relationship between different
types of mycorrhizae and to the factors that are the source of variability in types of
mycorrhizae.
ACKNOWLEDGMENTS
This study was supported by Cooperative Grant #60-PNW-79 with the Pacific
Northwest Range and Experiment Station, Portland, Oregon and by McIntyre-Stennis
Grant #48 Humboldt State University, Arcata, California.
REFERENCES
Comes, M. 1980. Some mycorrhizal syntheses on Arctostaphylos manzanita, Arbutus men-
ziesti and selected conifers. Masters’ Thesis, Humboldt State University.
Harley, J.L. 1959. Biology of mycorrhizae. Interscience Publishers, Inc. New York.
Largent, D.L., N. Sugihara, and A. Brinitzer. 1980a. Amanita gemmata, a non-host-
specific mycorrhizal fungus of Arctostaphylos manzanita. Mycologia 72: 435-39.
Largent, D.L., N. Sugihara, and C. Wishner. 1980b. Mycorrhizae of ericaceous shrubs and
subshrubs in Northern California. Can. J. Bot. 58: 2274-79.
Majstrik, V.K. and E. Hadac. 1975. Mycorrhizas of Arctostaphylos uva-ursi. Pedobiologia
15(5): 336-42.
Molina, R. and J.M. Trappe. 1982. Lack of mycorrhizal specificity in the ericaceous hosts
Arbutus menziesii and Arctostaphylos uva-ursi. New Phytol. 90/3. (In Press)
Trappe, J.M. 1964. Mycorrhizal hosts and distribution of Cenococcum graniforme. Lloydia
27: 100-06.
Zak, B. 19738. Classification of ectomycorrhizae. pp 48-78. In: Ectomycorrhizae; their ecol-
ogy and physiology. Ed. G.C. Marks. Academic Press, New York.
Zak, B. 1974. Ectendomycorrhizae of Pacific Madrone (Arbutus menziesii). Trans. Brit.
Mycol. Soc. 62: 210-14.
Zak, B. 1976a. Pure culture synthesis of bearberry mycorrhizae. Can. J. Bot. 54: 1297-
1305.
Zak, B. 1976b. Pure culture synthesis of Pacific madrone ectendomycorrhizae. Mycol. 63:
362-69.
MYCOTAXON
VoOlOAVI,, No, 25. pp. 537-556 January-March 1983
Fa nn lee eee ea
CHECK-LIST OF ROMANIAN PERONOSPORALES
O. CONSTANTINESCU* and G. NEGREAN
Instttutul de Sttinte Btologice, Herbarul micologic
Spl. Independentet 296, R-77748 Bucuregtt 17, Romanta
Together with the Uredinales, Ustilaginales and Erysipha-
les, the Peronosporales have been intensively studied in
Romania. The most valuable contributions are those by T.
bovulesct, IT. Rayss, C.V. Oescu, LL. Radulescu. and OV Ssavu-
lescu. Over the years other students have recorded many
downy mildews and T. & O. Savulescu have published a list
'Peronosporaceele din Republica Populara Roman&' (Lucr.
Grad. bot. Buc. 1963: 1-146, issued 1964) based on litera-
ture data. In a recent paper (Constantinescu & Negrean,
Feddes Repert. 93 (9/10) 1982, in press) 8 species and 177
fungus/host combinations were newly recorded for Romania,
and 80 plants not previously recorded as hosts were listed.
Both the mentioned papers include short surveys of the re-
search done in Romania on these fungi. The available data
show that 641 out of c. 3000 phanerogamic taxa of Romanian
flora are parasitized by downy mildews. Romania, a relati-
vely small territory, apparently has the richest known flo-
ra of Peronosporales.
Although our search through the literature concerning Ro-
manian Peronosporales (about 280 publications) has been
thourough, this check-list is only based on an examination
of over 5000 specimens of which we collected more than 3000
between 1957-1981, particularly between 1975-1980. We have
also collected and preserved new specimens for most of the
known fungus/host combination which were not supported by
voucher herbarium material. This work was part of a project
concerning 'Cryptogamic Flora of Romania'. Almost all of
*
Present address: Uppsala Untverstty, Instttute of Systema-
tie Botany, P.O.Box 541, S-751 21 Uppsala, Sweden.
538
the material studied is preserved in the herbarium BUCM and
some specimens are deposited in BUC, CL, CRAF, I, IBIR and
SIB herbaria.
This list comprises only Albuginaceae and Peronosporaceae,
Pythtum and Phytophthora (Pythiaceae) were not included be-
cause they are little known in Romania. For these two gene-
ra only a limited number of herbarium specimens and living
cultures are available.
The main difficulty encountered during our study was the
absence of a modern, monographic treatment of such important
genera as Peronospora and Plasmopara. Consequently, we had
to take some taxonomical decisions, particularly in Perono-
spora, which are not in accordance with most of the publi-
shed studies. In general, we have recognized only those dow-
ny mildew species within a host family, which can be delimi-
ted by qualitative morphological features e.g. shape of co-
nidia or sporangia (length/width ratio), oospore ornamenta-
tion, tip of ultimate branchlets, and not by quantitative
characters such as the size of conidia or other organs.
However, in the treatment of Peronospora parasitic on some
host families (Boraginaceae, Caryophyllaceae, Leguminosae)
this principle could not be applied consistently. The oospo-
re ornamentation of Peronospora inhabiting these plants
proved to have diagnostic value but in many micro-species
no oospores could be found. Consequently, they were partly
treated in a conservative way. These decisions can be ques-
tioned, particularly because they are not based on the study
of the type specimens of taxa concerned and, actually, we
changed a few times our mind during the final part of our
work.
The physiological specialization below host family level,
extensively used in the past but only rarely properly pro-
ved, was not taken into account for species delimitation.
No infraspecific taxa are recognized in this paper and no
'formae speciales' are mentioned, though the latter are
often used by plant pathologists.
As a result of a broader species concept that the one
adopted by T. & O. SA&vulescu (1.c.), the number of 'good'
species of Peronosporales was reduced from 254 to 112, al-
though the number of hosts has increased by c. 274 taxa
Cabte: B)
In a few instances authors are cited according to the
changed Art. 13 ICBN adopted by the Sydney Bot. Congr.
(Taxon 30: 106, 905. 1981). Author's names are abbreviated
Doo
Table 1
SYNOPTIC TABLE OF ROMANIAN PERONOSPORALES
CENEBRA number _ humber of hosts
of fami- species &
, ; genera
spectres bres subsp.
Albugo 6 ig 60 141
Bastdtophora 1 1 1 1
Bremta a 1 23 82
Bremtella i 1 | &)
Peronospora 88 oe 17 438
Plasmopara 10 9 Sih Dit
Pseudoperonospora 4 y 4 ih
Selerospora Hl 1 1 3
TOTAL
Shun gi TD = = a
- fungus/host comb. ~ 56 286 Se
- hosts - he eat 641
according to Hawksworth (Rev. Pl. Pathol. 59: 473-480. 1980).
The nomenclature of the hosts is in accordance with 'Flora
Europaea'. Genera and species are arranged alphabetically,
except for Peronospora species which are listed according
to alphabetically arranged host families; a host-fungus
index is added. The first author is responsible for the
identification, taxonomy and nomenclature of the fungi and
the second author for the same with regard to the host
plants.
We are grateful to I. Comes (CRAF), M. Hatmanu (IBIR), G.
Nedelcu (BUC), 0. Ratiu (CL), E. Schneider (SIB) and C. Sir-
bu (I) for the loan of specimens in their keeping, and to
W. Gams, CBS Baarn, for redactional help.
540
ALBUGO (Pers.) Roussel
A. bliti (BivJ-—Bern.) Kuntze:.on Amaranthus valbus. i. Aveda
toides 6. Watson. Al? bouchonii Thelin AS *eraeci Zancai
AY hybridus b..thy lividus Ls. A. wretrofhexusaG.
A. candida (J.F. Gmelin : Fr.) Kuntze on Alyssum alyssoides
(L.) L., A. borzeanum E.I. Nya@rAdy, A. caliacrae E.I. Nyara-
dy, A. desertorum Stapf, A. hirsutum Bieb.,, A. .linwiel sum
Stephan ex Willd., A. minutum Schlecht. ex DC., A. murale
Waldst. & Kit., 9A. petraeum Ard., A. saxvatile Lege Ae ator.
osum Willd.; Arabidopsis thaliana (L.) Heynh.; Arabis alpina
Ls, AL glabra (i. Bernh. 5A. Hirsute: (iv) Scop.) As mee
Bertol.; Av procurrens Waldst.. & Kit. 2s uAt erecta. (i ee
sagittata (Bertol.) DC., A. turrita L.; Armoracia rusticana
P. Gaertner, B. Meyer & Scherb.; Barbarea vulgaris R. Bru;
Berteroa incana (L.)DC.; Biscutella laevigata L.; Brassica
elongata Ehrh. subsp. elongata; B. juncea (L.) Czern. &
Coss... Bi mapus le (cult), Be niera sie) Koch bp iecoben es
bsg ba rena @lns Cand ee Be rapa i. subst. Ole teraspo.
(cult.); Bunias orientalis L.; Calepina irregularis (Asso)
Thell.; Camelina microcarpa Andrz. ex DC., C. rumelica Ve-
lien;, ¢@. sative (G.) Crantz: Capsella -bursa=pastoris (my
Medicus Cardamine amara h., Cy hirsuta i, Ce metus
Moretti, C. pratensis L.; Cardaminopsis arenosa (L.) Hayek,
C. halleri. (L.) Hayek subsp. halleri, C. halleri subspaiovi=
rensis (Wulfen) Hegi & E. Schmidt, C. neglecta (Schultes)
Hayek; Cardaria draba (L.) Desv.; Cheiranthus cheiri L.
(cult. ); Chorispora tenella (Pallas) DC.; Cleome spinosa
Jacq. (cult. ); Conringia orientalis (la) Dumortia; Coroncpue
squamatus (Forsk&al) Ascherson; Descurainia sophia (L.) Webb
ex Prantl; Diplotaxis muralis (L.) DC.; Draba lasiocarpa
Rochel (incl. subsp. elongata (Host) Jav.); Erophila verna
(L.) Chevall. subsp. verna; Erysimum cheiranthoides L., E.
comatum Pantit, E. crepidifolium Reichenb., E. cuspidatum
(Bieb.) “DC., EB. diffusum Ehrh., EH: odoratum Ehrh., B..repans
dum L., E. witmannii Zawadski; Euclidium syriacum (L.) R.
Br.; Hesperis matronalis (cult.); Hornungia petraea (L. )
Reichenb.; Hutchinsia alpina (L.) R. Br. subsp. brevicaulis
(Hoppe) Arcangeli; Hymenolobus procumbens (L.) Nutt. ex Tor-
rey & A. Gray; Iberis saxatilis L., I. umbelkata i. 3; Leatis
tinctoria L.; Lepidium campestre (L.) R. Br., Ly, cartilagi—-
neum (J. Mayer) Thell. subsp. crassifolium (Waldst. & Kit.)
Thell..) UG. censiftiorum Schrader, L. Latafoliumal, tapers
liatum L., L. ruderale L.. Lunaria annua L. subsp. pachyr-
rhiza (Borbas) Hayek; Neslia paniculata (L.) Desv.; Rapha-
541
nus raphaniastrum L. subsp. raphaniastrum, R. sativus L.
(cult.); Rorippa amphibia (L.) Besser, R. austriaca (Crantz)
Besser, R. islandica (Oeder) Borb&as, R. pyrenaica (Lam.)
Reichenb., R. sylvestris (L.) Besser subsp. sylvestris, R.
sylvestris subsp. kerneri (Menyh.) Sod; Sinapis alba L.
(cult.), S. arvensis L.; Sisymbrium altissimum L., S. loese-
(ime. os Orlicinale’ (i.) scopes oO. Orlentale In, so. poly—
morphum (Murray) Roth; Syrenia cana (Piller & Mitterp. )
Neilr., S. montana (Pallas) Klokov; Thlaspi arvense L.; T.
perfoliatum L.
A. caryophyllacearum (Wallr.) Cif. & Biga on Spergularia
mamiGesile) Griseb. (= S. salanad. &.C. Presi.) 76. media
(bwec. Presi -(=.S. marginata Kittel).
A. eurotiae Tranzschel on Ceratocarpus arenarius L.
A. portulacae (DC.) Kuntze on Portulaca eleracea (U.iisubsp.
oleracea, P. oleracea subsp. sativa (Haw.) Celak. (cult.)
A. tragopogonis (Pers.) Gray on Ambrosia artemisiifolia L.;
Artemisia vulgaris L.; Centaurea atropurpurea Waldst. &
here Kouschyana Heurtrel,,'C. orientalis ..7C. ‘7 scabio—
sa L.; Chamomilla suaveolens (Pursh) Rydb.; Cirsium alatum
(S.G. Gmelin) Bobrov (= C. setigerum Ledeb.), C. arvense
(ey cop., ©. canum (Ty. ) Scop. ; C. erisithales ‘((aca. }
PCOy. nc. Oleraceum (L.) Scop... C. rivulare (Jacq) AIT S,
C. vulgare (Savi) Ten.; Crepis sancta (L.) Babcock (= Lago-
Seris sancta (L.) K. Mal¥); Crupina vulgaris Cass.; Filago
VouLearicrlam.; Inula britannica: L., I) conyza De: , F. ‘sali=
cina L.; Logfia arvensis (L.) Holub; Scorzonera cana (C.A.
Meyer) O.Hoffm. S. hispanica L., S. laciniata b.; Senecio
vernalis Waldst. & Kit.; Serratula tinctoria L.; Tragopogon
dubeuewocoo sh. porrifolius i, cult.) . Te pravensis Leics
1.3; Xeranthemum annuum L., X. cylindraceum Sibth. & Sm.
BASIDIOPHORA Roze & Cornu
B. entospora Roze & Cornu on Conyza canadensis (L.) Cronq.
(= Erigeron canadensis L.).
BREMIA Regel
B. lactucae Regel on Arctium lappa L., A. minus Bernh., A.
tomentosum Miller; Carduus acanthoides L., C. crispus L..
Penibanceiia. Ceiperconate (1, ),.Jacd., S.L. subsp: albacans
(Adamovié) Kazmi; Carlina acaulis L., s.l.; Centaurea cya-
Busiise, 6, gacea L., C. mollis Waldst. & Kit.; C. migres—
542
cens Willd., C. pannonica (Heuffel) Simonkai subsp. pannoni-
ca\, (C.. phrygia LL. subsp. phryria, C. phrygia subep. caspse
tica (Porcius) Dostal, C. phrygia subsp. melanocalathia
(Borbas) Dostal, C. phrygia subsp. pseudophrygia (C.A. Me-
yer) Gugler, C. rocheliana (Heuffel) Dostal (= C. banatica
Rochel ex Hayek, non A. Kerner), C. solstitialis L., s.l.,
C. stenolepis A. Kerner subsp. stenolepis, C. triumfetti
All.; Cephalorrhynchus tuberosus (Steven) Schschian; Cicer-
bita alpina (L.) Wallr.; Cirsium arvense (L.) Scop., Co ca=
num “(L.) AlTs5 C.- oleraceum (L.)) Scop., (Co palustre (ie
Scop., Ci. rivulare’ (Jacqg.) Alls, (C. wolgare (Savi) Vea
Crepis biennis L., C. foetida L. subsp. rhoeadifolia (Bieb.)
Celak. : C. paludosa (L.) Moench, C. pulchra L., C. sancta
(L.) Babcock (= Lagoseris sancta (L.) K. Mal¥), C. setosa
Haller fil.; Dimorphotheca pluvialis Moench (cult.); Gazania
Yigens (L.) Gaertner (= G. splendens E.G. & A. Henderson);
Helichrysum bracteatum (Vent.) Andrews (cult.); Hieracium
argillaceum' Jordan, 's.1., H. ‘aurantiacum Us, H. bifidum
Kit., H. caesium (Fries) Fries, H. caespitosum Dumort., H.
eymosum.L.-, HH. % Lactucella Wallr.,.H. Jactucella Walia
subsp. lactucella, H. murorum L., H. pilosella; Ll. subsp...
losella, H. pilosella subsp. micradenium Negeli & Peter, H.
pliloselloides Vill. subsp. piloselloides, H. praealtum Vill.
ex Gochant, s.1., H. praealtum subsp. bauhinii (Besser) Pe-
tunnikov, H. praealtum subsp. thaumasium (Peter) P.D. Sell,
H. rotundatum Kit. & Schultes (= H. transsilvanicum Heuf-
fel), H. umbellatum L.; Hypochoeris radicata L.; Lactuca sa-
ligna Ivy L. sativa Le (cult. ) 2 U2 serriolay i. ,ile wamines
(L.) J. & C. Presl; Lapsana communis L., L. intermedia
Bieb.;. Leontodon, autumnalis L., L. hispidus Lajcs-l) ace.
lis muralis (L.) Dumort.; Onopordum acanthium L., 0. tauri-
cum Willd.; Picris echioides L. (= Helminthia echioides (L.)
Gaertner), P. hieracioides L., s.1.; Senecio aquaticus Hill
subsp. barbareifolius (Wimmer & Grab.) Walters, S. squalidus
Li. (= "S. ‘rupestris Waldst: & Kite), 8S. subalpinus Keen soe
vernalis Waldst. & Kit.,) S. vulgaris L.i; Sonchus arvensis
l.5 S-1., S. arvensis subsp. wliginosus (Bieb.) Nyman o.
asper (L.) Hill subsp. asper, S. asper subsp. glaucescens
(Jordan) Ball; S. oleraceus L., S. palustris b., Taraxacum
‘officinale Weber'; Xeranthemum annuum L.
BREMIELLA G. Wilson
B. baudysii (Skalicky¥) O. Const. & Negrean on Berula erecta
(Hudson) Coville (= Sium erectum Hudson); Oenanthe silaifo-
Ina Brebl:?Sium Jatifolaum ty.
543
PERONOSPORA Corda
on BORAGINACEAE
P. asnperuginis Schrdéter on Asperugo procumbens L.
P. cerinthe Uliyanishchev ex Jacz. & P.A. Jacz. on Cerinthe
minor L.
P. cynoglossi Burrill ex Swingle on Cynoglossum hungaricum
Simonkai.
P. echii (L. Krieger) Jacz. & P.A. Jacz. on Echium vulgare
Le
P. echinospermi (Swingle) Swingle on Lappula barbata (Bieb.)
Girke, L. marginata (Bieb.) Glirke (= L. patula (Lehm.) Me-
nyh.), L. squarrosa (Retz.) Dumort. subsp. squarrosa (= L.
echinata Fritsch).
P. lithospermi G&um. on Buglossoides arvensis (L.) I.M.
Johnston subsp. arvensis (= Lithospermum arvense L.), B. ar-
vensis subsp. sibthorpiana (Griseb.) R. Fernandes, B. glan-
dulosus (Velen.) R. Fernandes (= Lithospermum glandulosum
Velen. ).
P. myosotidis de Bary on Myosotis alpestris F.W. Schmidt
(cult.), M. arvensis (L.) Hill subsp. arvensis, M. discolor
Pers. subsp. discolor (= M. versicolor Sm.), M. ramosissima
Rochel subsp. ramosissima, M. scorpioides L. (= M. palustris
(L.) Hill), M. sparsiflora Mikan ex Pohl, M. stricta Link ex
Roemer & Schultes, M. sylvatica Hoffm subsp. sylvatica.
P. rocheliae Kalimbetov on Rochelia disperma (L. fil.) C.
Koch subsp. retorta (Pallas) E. Kotejowa.
P. symphyti Gaum. on Symphytum cordatum Waldst. & Kit., S.
mojosimeochnur, GS. officinale l.,°s.1.,,S. officinale subsp.
uliginosum (A. Kerner) Nyman, S. ottomanum Friv.
on CAMPANULACEAE
P. phyteumatis Fuckel on Phyteuma vagneri A. Kerner.
on CAPPARIDACEAE
P. parasitica (Pers. : Fr.) Fr. on Cleome spinosa Jacq.
bend ta)
on CARYOPHYLLACEAE
P. agrostemmatis Gdum. on Agrostemma githago L.
P. alsinearum Casp. on Stellaria media (L.) Vill.
544
P. arenariae (Berk.) Tul. on Moehringia trinervia (L.)
Chatxv.
P. campestris Gaum. on Arenaria serpyllifolia L.
P. conferta (Unger) Unger on Cerastium dubium (Bast.) 0. Sch-
warz (= C. anomalum Waldst. & Kit.), C. fontanum Baumg. sub-
Spentriviale (link) Jalas:
gypsophilae Jacz. & P.A. Jacz. on Gypsophyla muralis L.
holostei Casp. ex de Bary on Holosteum umbellatum L., s.
., H. umbellatum subsp. glutinosum (Bieb.) Nyman.
P
P
i
P. lychnitis Gaum. on Lychnis alpina L. (cults), Lb. visearae
iy.
P
melandrii Gaum. on Silene alba (Miller) E.H.L. Krause, S.
noetifiora ley Selotites (le )owaber.
P. obovata Bonorden on Spergula arvensis L.; Spergularia ma-
rina {(l.) Griseb.,°S. media (L..) °C. Pres, S:¥rubra (ee
eC. Presi,
P. parva Gaum. on Stellaria holostea L.
P, scleranthi Rabenh. on Scleranthus annuus L., s.1.3°S 7a
eivatus .ochurs
P. tomentosa Fuckel on Cerastium arvense L., s.1.(cult.)jVG3
brachypetalum Pers., C. glomeratum Thuill., C. semidecandrum
Ins
on CHENOPODIACEAE
P. chenopodii-glauci Gaum. on Chenopodium glaucum L.
P. farinosa (Fr.) Fr. on Atriplex hastata L.,; AY hortensae
L. (ewlte.), A. mitens Schkuhr. A. oblongifolia Walder.
Kit., A. patula L., A. rosea L., A. tatarica L.; Beta vues
ris L. (cult.); Ceratocarpus arenarius L.; Chenopodium album |
L.,. S.l.«,, C. ambrosioides L., C. bonus—henricus. L. 5 C.oiae
folium las, C. hybridum L., C..murale Lec. opi comm
Schrader ex Koch & Ziz, C. polyspermum L., C. urbicum) lL. jee
vulvaria L.; Halimione verrucifera (Bieb.) Aellen (= Obione
verrucifera (Bieb.) Moqg.); Kochia scoparia (L.) Schrader
(cult.J= Spimnacia oleracea L.w(eult.)
P. vistulensis Wrébl. on Corispermum nitidum Kit.; Salsola
kali L.,.c) Kali L. subsp. ruthenica 115 1m eeoo, |
on CISTACEAE
P. leptoclada Sacc. on Helianthemum canum (L.) Baumg. subsp.
545
canum, H. nummularium (L.) Miller, s.1., H. nummularium sub-
sp. nummularium, H. nummularium subsp. obscurum (Celak.) J.
Holub, H, ‘oclendicum (1. DC. subsp, alpestre (Jacq. )) Breie
Sureeetl, salicifolium (i. Miller’
on COMPOSITAE
P. leptosperma de Bary on Anthemis arvensis L., A. austria-
CanJecq. , A. cotula L.o;-Artemisia absinthium Liv,’ A.” annua
L.; Chamomilla recutita (L.) Rauschert, C. suaveolens
(Pursh) Rydb.; Leucanthemum nipponicum Franch. (= Chrysan-
themum nipponicum Matsum.) (cult.); Matricaria perforata
Mérat (= M. inodora Lee). Tanacetum vulgare L.
P. radii de Bary on Anthemis ruthenica Bieb.; Matricaria
perforata Mérat.
on CRUCIFERAE
P. lepidii (McAlp.) G. Wilson on Coronopus squamatus (For-
sk&4l) Ascherson; Lepidium densiflorum Schrader, L. ruderale
i.
P. niessleana Berl. on Alliaria petiolata (Bieb.) Cavara &
Grande.
P. parasitica (Pers. : Fr.) Fr. on Alyssum alyssoides (L.)
is os Cesertorum Stapt, A. hirsutum Bieb., A. Dinifolium
Stephan ex Willd., A. minutum Schlecht. ex DC., A. murale
Waldst. & Kit., A. petraeum Ard., A. saxatile L.; Arabidop-
Sis thaliana (L.) Heynh.; Arabis alpina L., A. glabra (L.)
Rerun, 4. hirsuta (i.) Scop. A. turrita l.;)Armoracia
rusticana P. Gaertner, B. Meyer & Scherb.; Barbarea vulga-
ris R. Br.; Berteroa incana (L.) DC., Brassica elongata
EBhrh. subsp. elongata, B. juncea (L.) Czern. & Coss., B. na-
Posies eult..) UB. niera (Is) Koch, B. oleraccaule, inel,
Joy merapa In .-S.L., B. rapa, subsp..olei tera, DC. (culty)
Bunias orientalis L.; Calepina irregularia (Asso) Thell.;
Panelina microcarpa Andrz. ex DC., C.. rumelica Velen., Cc.
Sativa (L.) Crantz; Capsella bursa-pastoris (L.) Medicus;
Cardamine bulbifera (L.) Crantz (= Dentaria bulbifera L.),
ce. flexuosa With., C. glanduligera O. Schwarz (= Dentaria
plarditosa Waldst.& Kit.) >) C. graeca lo, Cy hirsuta. Lin, Ce
impatiens L., C. matthiolii Moretti; Cardaminopsis arenosa
(L.) Hayek; Cardaria draba (L.) Desv. subsp. draba; Cheiran-
ehus cheira i. (cult. ); Chorispora tenellas(pallas) DCs, Co-
chlearia pyrenaica DC., var. borzeana Coman & E.I. Nyarady,
Conringia orientalis (L.) Dumort.; Descurainia sophia (L.)
Webb ex Prantl (= Sisymbrium sophia L.); Diplotaxis muralis
L., D. tenuifolia (L.) DC.; Draba nemorosa L.; Erophila ver-
546
na (L.) Chevall., s.1l.; Erucastrum nasturtiifolia (Poiret)
O.E. Schulz; Erysimum cheiranthoides L., E. cuspidatum
(Bieb. ) Dc. (= Syrenia cuspidata L.); EB. diftfausum Shriewe
odoratum Ehrh., BE. repandum ;, E. witmannii Zawadski, sviw-
Euclidium syriacum (L.) R. Br.; Hesperis matronalis L., s.
1., H. sylvestris Crantz subsp. velenovskyi (Fritsch) Borza;
Iberis umbellata L.; Isatis tinctoria L., s.1.3 Lepidium
campestre (L..) R.. Br., L. densiflorum Schrader, L. 1aGiros
lium L., L. perfoliatum L.;/Lobularia maritima, (i). Dec
(cult.); Lunaria annua L. subsp. pachyrrhiza (Borb&as) Hayek,
L. ‘rediviva L.; Matthiola incana (L..) Ry Br. subsp.) inecana
(cult.), M. longipetala (Vent...) DC. subsp. bicornisy(oipome
& Sm.) P.W. Ball; Nasturtium officinale R. Br.; Neslia pani-
culata (L:) Desv., s.1.; Peltaria alliacea Jacq.; Raphanus
raphaniastrum L. subsp. raphaniastrum, R. raphaniastrum sub-
sp. landra (Moretti ex DC.) Bonnier & Layens, R. sativus L.
(cult.); Rapistrum perenne (L.) All.; Rorippa amphibia (L.)
Besser, R. x armoracioides (Tausch) Fuss, R. austriaca
(Crantz) Besser, R. islandica (Oeder) Borbas, R. sylvestris
(L.) Besser subsp. sylvestris; Sinapis alba L. (eult.) jis.
arvensis L.; Sisymbrium altissimum b., S. loeselia; bie.
officinale (L.) Scop., S. orientale L., S. polymorphum (Mur-
ray) Roth, S. strictissimum L.; Syrenia montana (Pallas)
Klokov; Thlaspi arvense L., T. perfoliatum L.
on DIPSACACEAE
P. cephalariae-laevigatae Savul. & Rayss on Cephalaria lae-
vigata (Waldst. & Kit.) Schrader, C. uralensis (J. Murray)
Schrader subsp. multifida (Roman) Roman & Beldie.
P. dipsaci Tul. ex de Bary on Dipsacum fullonum L. (= D.
sylvestris Hudson), D. laciniatus L., D. sativus (L.) Hon-
ckeny (cult.).
P. knautiae Fuckel ex Schréter on Cephalaria transylvanica
(L.) Roemer & Schultes; Knautia arvensis (L.) Coulter, K.
? dipsacifolia Kreutzer s.1., K. drymeia Heuffel subsp. dry-
meia; Scabiosa columbaria L. subsp. columbaria (="S...bana tas
ea. Waldst.*tekite)> S. lucida Vill. peubsp. eeiaar.
echroleuca Uc
P. violacea Berk. & Cooke on Knautia arvensis (L.) Coulter
(in flowers); Scabiosa columbaria L. (in flowers).
on EUPHORBIACEAE
P. cyparisiae de Bary on Euphorbia amygdaloides L., E. cypa-_
rissias L., E. esula L. subsp. tommasiniana (Bertol.) Nyman
(= BE, vaegeta auct..).
Constantinescu & Negrean 9
547
P. euphorbiae Fuckel on Euphorbia ? serrulata Thuill.
P. valesiaca Gdum. on Euphorbia seguierana Necker (= E. ge-
rardiana Jacq.).
on GENTIANACEAE
P, erythraeae Kuhn ex Gadum. on Centaurium erythraea Rafn
subsp. erythraea, C. pulchellum (Swartz) Druce.
on GERANIACEAE
P. conglomerata Fuckel on Erodium ciconium (L.) L'Hér., E.
cicutarium (L.) L'Hér., E. hoefftianum C.A. Meyer (incl. E.
neilreichii Janka); Geranium columbinum L., G. dissectum L.,
Gaeucidum ls G.;molle L., G: phaeum L., G. pusillum L.,
Vancopercianum. Ls... sylvaticumsL.
on LABIATAE
P. lamii A. Braun on Acinos alpinus (L.) Moench subsp. alpi-
nus, A. alpinus subsp. majoranifolius (Miller) P.W. Ball,
A. arvensis (Lam.) Dandy; Lamium album L., L. amplexicaule
L., L. garganicum L. subsp. laevigatum Arcangeli (= L. bi-
thynicum Bentham), L. maculatum L., L. purpureum L.; Leonu-
rus marrubiastrum 1.3; Stachys palustris L.; Ziziphora capi-
tata L.
P. stigmaticola Raunkiaer on Mentha longifolia (L.) Hudson
(in flowers).
on LEGUMINOSAE
P. aestivalis H. Sydow in Gaum. on Medicago falcata L., M.
sativa L. (cult.); Melilotus alba Medicus, M. officinalis
(L.) Pallas.
P. astragalina H. Sydow on Astragalus cicer L., A. hamosus
eee onoprychis L.
P. coronillae Gaéum. on Coronilla scorpioides (L.) Koch, C.
vari4 Ls
Peeeryvs) As Gust. ,on Vicia. mirsuta (L.) S.FisGray.
P. lotorum H. Sydow on Lotus tenuis Waldst. & Kit. ex Willd.
P. manshurica (Naum.) H. Sydow on Glycine max (L.) Merr.
Powlts).
P. medicaginis-minimae Gaponenko on Medicago lupulina L., M.
S minima (L.) Bartal.
P. senneniana Frag. & Sacc. on Lathyrus niger (L.) Bernh.
Bubpdu.ttcer. Uy. palustris bL., L. tuberosus i.
P. tetragonolobi Gdum. on Tetragonolobus maritimus (L.) Roth
(=P esilaquosucse( ua Roth).
P. trifoliorum de Bary on Trifolium alpestre L., T. arvense
Ls, Ts. badium L., UT. campestre Schreber, T. dubaum Sipe
T. echinatum Bieb., T. hybridum L., T. hybridum subsp. ele-
gans (Savi) Ascherson & Graebner, T. medium L., s.1., T. me=
dium subsp. banaticum (Heuffel) Hendrych, T. montanum L., T.
pratense L.. (cult.), T. repens L., T. spadiceum L., etre
atum L.
P. ‘viciae (Berk. ) Casp. on Galega officinalis L.; Lathyrus
eicera L.; LL. bhirsucus’L., lL. nissolia Ge, Le tspnseraer.
Retz. , Le sylvestris Le; Pisum sativum Le subsp. “sacar
(cult.); Vicia angustifolia L., V. cracca L., V. grandifolia
ecop.,; V. Lathyroides L., V. lutea L: subsp. Duteay Vue ce
bonensis*’L., V. pannonica Crantz, s.l., Ve -pannoniesg-u
striata (Bieb.) Nyman, V. peregrina L., V. pisiformis saa
sativa L. (cults), V. sepium Li,)V. tetrasperma lo. Vee
losa/Roth.
on LILIACEAE
P. destructor (Berk. ) Casp. in Berk. on Allium cepa L.
(cubis Pistulesum sine (emltauns
on LINACEAE
P. lini Schrdter on Linum catharticum L.
on PAPAVERACEAE
P.affinis Rossm. on Fumaris officinalis L., F. ischieieners
Soyer-Willemet, F. vaillantii Loisel.
P. arborescens (Berk.) Casp. on Papaver dubium L., P. rhoeas
Ea esomnat erum We (cule)
P. corydalis de Bary on Corydalis bulbosa (L.) DC. subsp.
Marschalliana (Pallas) Chater, C. solida (L.) Schwarz; Gla-
ucium corniculatum (L.) J.H. Rudolph.
P. cristata Tranzschel on Papaver hybridum L.
P. corydalis-intermediae Gdum. on Corydalis capnoides (L.)
ers.
on PLANTAGINACEAE
P. alta Fuckel on Plantago altissima bL., P..cornutz Gouan.
P. lanceolata L.,.P. major L....ssl., P. major ssubsp a eemes
media (DC) Arcangeli, P. media L., s.1.
P. plataginis Burrill on P. arenaria Waldst. & Kit. C= ps
549
acca Lies), Po» -Major Ie. GS. dre
on PLUMBAGINACEAE
P. statices Lobik on Limonium gmelinii (Willd. ) Kuntze, L.
vulgare Miller subsp. serotinum (Reichenb.) Gams (= Statice
limonium L. )
on POLYGONACEAE
P. ducometii Siem. & Jank. on Fagopyrum esculentum Moench
Gell ea:)
P. polygoni Thi. ex A. Fischer on Polygonum arenarium Wald-
Stu. Kit. subsp. arenarium, P. aviculare: Ll.
P. polygoni-convolvuli A. Gust. on Bilderdykia convolvulus
(i) Damort .
P. yuimers) Corda on Rumex acetosa I. ; R. acetosella' Ly, R.
alipestris Jacq. (= Ra arifolius All.); R. tuberesus Lb. sub-
Sp... tuberosus .
on PRIMULACEAE
P. androsaces Niessl on Androsace elongata L.; Primula veris
in
on RANUNCULACEAE
P. ficariae Tul. ex de Bary on Helleborus purpurascens Wald-
pe woeekic.:) Ranunculus. acrig Li. Snbsp. “acris, Rh. acris sub—
sp. strigulosus (Schur) Hyl. (= R. stevenii auct.), R. bul-
bosis f. subsp... bulbosus, R- cassubicus &.5 Rs carpaticus
Herbich, R. constantinopolitanus (DC.) D'Urv., R. crenatus
Vero ambi he. ficaria 4. ,' he tLammuba ie. 6.1. , h.: a=
werigvorus DC., KR: oreophilus Bileb., R. oxyspermus Willd,
Rerpedauus Waldst. & Kit.5, RK. phatanifolius G.. RR. polyan—
themos L. subsp. polyanthemos, R. pseudomontanus Schur, R.
Tenens din. Khwtsardous Crantz.
Preetli¥rica Gaum. On Ranunculus iliyricus L:
P. parvula A. Schneider ex Jacz. & P.A. Jacz. on Isopyrum
thalictroides L.
on RESEDACEAE
P. crispula Fuckel on Reseda lutea L., R. phyteuma L.
on ROSACEAE
P. potentillae de Bary on Potentilla aurea L. subsp. aurea,
P. aurea L. subsp. chrysocraspeda (Lehm.) Nyman (= P. terna-
ra wn Koch )'.
550
P. sparsa Berk. ex Berk & Broome on Alchemilla glabra Ney-
gent.’, A. “hyborida” suct., A. "“vulgaris’ auction
chlora Rothm.; Aremonia agrimonoides (L.) DC. subsp. agrimo-
noides; Fragaria vesca L.; Geum aleppicum Jacq., G. coccine-
um Sibth. & Sm. (cult.), G. rivale L., G. urbanum L.: Poten=
tills ansering) l.. iP. reptans,L.; Rosa ycaliaica lace ube
caesius L. 5 Revcanescens DC., R. hirsutus Waldst. Kee wee
idaeus L.; Sanguisorba minor Scop.
on RUBIACEAE
P. calotheca de Bary on Asperula purpurea (L.) Ehrend. sub-
Sp. purpurea (= Galium purpureum L.); Cruciata pedemontana
(Bellardi) Ehrend. (= Galium pedemontanum (Bellardi) All.).;
Galium album Miller, G. aparine L., G. boreale L., G. diva-
ricatum Pourret ex Lam., G. humifusum Bieb., Gy. mollveow.
G. odoratum Scop. (= Asperula odorata L.), G. palustre L.,
G. rubioides L,))'G. schultesii Vest.)°G. spurituw i.09%G. oe
cornutum Dandy, Gt verum L. subsp. verum:
P. sherardiae Fuckel on Sherardia arvensis L.
on SAXIFRAGACEAE
P. chrysosplenii Fuckel on Chrysosplenium alternifolium L.
on SCROPHULARIACEAE
P. grisea (Unger) Unger on Chaenorrhinum minus (L.) Lange
subsp. minus; Digitalis grandiflora Miller; Euphrasia pecti-
nata Ten.; Linaria genistifolia (L.) Miller subsp. genisti-—
folia, L. vulgaris Miller; Veronica anagallis-aquatica L.,
V. anagalloides Guss., V. becccabunga L., V. catenata Pen-
nell (= V¥. aquatica Bernh., non .S.F. Gray), V..cftlcimamae
L., V. scardica Griseb., V. seutellatashL., V. serpy llitolee
L., V. serpyllifolia subsp. humifusa (Dickson) Syme, V. spi-
Cata lo subsp. spicata, Ve urticifolia Jacq.
P. melampyri (Bucholtz) J. Davis on Linaria genistifolia
(L.) Miller subsp. genistifolia , L. vulgaris Miller (in
flowers); Malampyrum arvense L., M. cristatum L., M. saxosum
Baumg.; Rhinanthus rumelicus Velen.
P. sordida Berk. & Broome on Scrophularia nodosa L., S. sco-
polii Hoppe; Verbasum banaticum Schrader, V. blattaria L.
V. densiflorum Bertol. (= V. thapsiforme Schrader) (cult.),
V. glabratum Prib. subsp. glabratum, V. nigrum i... eee
phlomoides L., V. speciosum Schrader, V. thapsus L293 .025
V. thapsus subsp. crassifolium (Lam.) Murb. (= V. montanum
Schrader); Veronica arvensis L., V. austriaca L. subsp. teu-
crium (L.) DA, Webb (= V. téeucrium L.). V. chamaedrys: ta.
551
Suey. nederifolia Lip Ve persica Poiret, Vo polrtarrries
(= V. didyma auct., vix Ten.), V. praecox All., V. prostra-
Ce, ity,
on SOLANACEAE
P. hyoscyami de Bary on Hyoscyamus niger L.
P. tabacina Adam on cultivated Nicotiana alata Link & Otto,
N. ipomopsiflora Dun., N. langsdorfii Schrank, N. longiflo-
renee NN. NOodlt Lora Cav... N..paniculaca Ls, Nierustica b.,
Weesueaveolens Lehm., N.. tabacum L.
on URTICACEAE
P. debaryi E. Salmon & Ware on Urtica urens L.
on VALERIANACEAE
P, valerianae Trail on Valeriana officinalis L.
P. valerianellae Fuckel on Valerianella dentata Pollich, V.
lasiocarpa (Steven) Betcke, V. locusta (L.) Letterade, V.
Dumida(L.) DCs; V. rimosa Bast.
on VIOLACEAE
P. violae de Bary ex Schréter on Viola arvensis Murray, V.
Picagpeliana ochultes, iV. tricolor L.. subsp. tricolor.
on ZYGOPHYLLACEAE
P. tribulina Pass. on Tribulus terrestris L.
PLASMOPARA Schréter
P. densa (Rabenh.) Schrdter on Euphrasia rostkoviana Hayne,
Pate stricta D. Wollf ex J.F. Lehm.; Odontites verna
(Bellardi) Dumort., s.1.; Rhinanthus alpinus Baumg., R. an-
Sustifolius C.C. Gmelin, R. minor L., R. rumelicus Velen.,
R. wagneri Degen.
P. epilobii (Otth) Sacc. & Sydow on Epilobium hirsutum L.,
EK. palustre L., E. parviflorum Schreber, E. roseum Schreber,
E. tetragonum L. subsp. lamyi (F.W. Schultz) Nyman.
P
. halstedii (Farlow) Berl. & de Toni on Carthamus tinctori-
us L. (cult.); Dimorphotheca sinuata DC. (cult.); Helianthus
annuus L. (cult.); Xanthium italicum Moretti, X. strumarium
Ty,
P. obducens (Schréter) Schréter on Impatiens noli-tangere
L.
P. pusilla (de Bary) Schréter on Geranium pratense L., G.
SoZ
sylvaticum L.
P. pygmaea (Unger) Schréter on Anemone nemorosa L., A. ra-
nunculoides L.; Helleborus odorus Waldst. & Kit. subsp. odo-
rus , H. purpurascens Waldst. & Kit.; Isopyrum thalictroi-
des L.
P. skvortzovii Miura on Abutilon theophrasti Medicus.
P. sphaerosperma Savul. on Tragopogon dubius Scop.
P. umbelliferarum (Casp.) Schréter ex Wartenw. on Aegopodium
podagraria L.; Ammi majus L. (cult.); Angelica archangelica
L., A. sylvestris L.; Anthriscus nitida (Wahlenb.) Garcke,
A. sylvestris (L.) Hoffm.; Apium graveolens L. Vcult. j2mca-
cucalis platycarpos L.; Conium maculatum L.; Daucus carota
L. (cult.); Ferulago campestris (Besser) Grec.; Foeniculum
vulgare Miller (cult.); Laserpitium latifolium L.; Léevisti-
cum officinale Koch (cult.); Ligusticum mutellina (L.) Cran-
tz; Meum athamanticum Jacq,; Pastinaca sativa L., s.l.
(spont. & cult.); Petroselinum crispum (Miller) A.W. Hill
(cult.); Peucedanum alsaticum L., P. austriacum (Jacq.)
Koch, P. palustre (L.) Moench; Pimpinella major (L.) Hudson,
P; saxifraga L.; Sanicula europaea L.; Silaum silaus (L.)
Schinz & Thell.; Smyrnium perfoliatum L.
P. viticola (Berk. & M:A. Curtis) Berl. (& de Toni on) Vie
Vinifera L. subsp. vinifera (cult.)> Ve vinifera subsp eer
vestris (C.C. Gmelin) Hegi, V. vinifera x American species
Cenk 8
PSEUDOPERONOSPORA Rostovtsev
P. cannabina (Otth) Curzi on Cannabis ruderalis Janisch., C.
sativa i. (eult i).
cubensis (Berk. & M.A. Curtis) Rostovtsev on Cucumis melo
Coultale Caasativus ne anew)
humuli (Miyabe & Takahashi) G. Wilson on Humulus lupulus
(cult. & spont.).
mo) Serle)" “eal ae)
. urticae (Lib. ex Berk.) E. Salmon & Ware on Urtica dioi-
ca L., U. kioviensis Rogow.
SCLEROSPORA “Scnréter
S. graminicola (Sacc.) Schréter on Setaria italica (L.)
Beauv., S. verticillata (L.) Beauv., S. viridis (L.) Beauv.
Soe)
HOST-FUNGUS INDEX
Host trinomials refer to subspecies. Whenever consecutive host species
are parasitized by the same fungus, the fungal name is only mentioned
after the last host listed. The following abbreviations are used throu-
ghout for fungal generic and specific epithets: A. = Albugo; B. = Bre-
mia; Ba. = Bastdtophora; Bl. = Bremtella; P. = Peronospora; Pl. = Plas-
mopara; Ps. = Pseudoperonospora; S. = Sclerospora,; cand. = candtda;
lact. = lactucae; paras. = parastttca; trag. = tragopogonts; umbel. =
umbellt ferarum.
ABUTILON theophrasti: Pl. skvortzovtt. ACINOS alpinus alpinus, alpinus
majoranifolius, arvensis: P. lamtt. AEGOPODIUM podagraria: Pl. umbel.
AGROSTEMMA githago: P. agrostemmatts. ALCHEMILLA glabra, 'hybrida',
'vulgaris', xanthochlora: P. sparsa. ALLIARIA petiolata: P. ntessleana.
ALLIUM cepa, fistulosum: P. destructor. ALYSSUM alyssoides: A. cand.,
P, paras.; borzeanum, caliacrae: A. cand.; desertorum, hirsutum, lini-
folium, minutum, murale, petraeum, saxatile: A. cand., P. paras.; tor-
tuosum: A. cand. AMARANTHUS albus, blitoides, ? bouchonii, graecizans,
hybridus, lividus, retroflexus: A. blttzt. AMBROSIA artemisiifolia: A.
trag. AMMI majus: Pl. umbel. ANDROSACE elongata: P. androsaces. ANEMONE
nemorosa, ranunculoides: Pl. pygmaea. ANGELICA archangelica, sylves-
tris: Pl. uwmbel. ANTHEMIS arvensis, austriaca, cotula: P. leptosperma;
ruthenica: P. radtt. ANTHRISCUS nitida, sylvestris: PZ. wnbel. APIUM
graveolens: Pl. wnbel. ARABIDOPSIS thaliana: A. cand., P. paras. ARABIS
alpina, glabra, hirsuta: A. cand., P. paras.; muralis, procurrens, rec-
ta, sagittata: A. cand.; turrita: A. cand., P. paras. ARCTIUM lappa,
minus, tomentosum: B. lact. AREMONIA agrimonoides agrimonoides: P.
sparsa. ARENARIA serpyllifolia: P. campestrts. ARMORACIA rusticana: A.
eand., P. paras. ARTEMISIA absinthium, annua: P. leptosperma; vulgaris:
A. trag. ASPERUGO procumbens: P. asperugtnts. ASPERULA purpurea: P. ca-
lotheea. ASTRAGALUS cicer, hamosus, onobrychis: P. astragaltna. ATRI-
PLEX hastata, hortensis, nitens, oblongifolia, patula, rosea, tatarica:
PR. fartnosa. BARBAREA vulgaris: A. cand., P. paras. BERTEROA incana: A.
cand., P. paras. BERULA erecta: Bl. baudystt. BETA vulgaris: P. farino-
Sa. BILDERDYKIA convolvulus: P. polygont-convolvult. BISCUTELLA laevi-
gata: A. cand. BRASSICA elongata elongata, juncea, napus, nigra, olera-
cea, rapa, rapa oleifera: A. cand., P. paras. BUGLOSSOIDES arvensis ar-
vensis, arvensis sibthorpiana, glandulosus: P. ltthospermt. BUNIAS ori-
entalis: A. cand., P. paras. CALEPINA irregularis: A. cand., P. paras.
CAMELINA microcarpa, rumelica, sativa: A. cand., P. paras. CANNABIS ru-
deralis, sativa: Ps. cannabtna. CAPSELLA bursa-pastoris: A. cand., P.
paras. CARDAMINE amara: A. cand.; bulbifera, flexuosa, glanduligera,
graeca: -P. paras.; hirsuta: A. cand., P. paras.; impatiens: P. paras.;
matthiolli: A. cand., P. paras.; pratensis: A. cand. CARDAMINOPSIS are-
nosa:,4. cand. , P:.paras.; haltTeri halleri,, halleri ovirensis, neglec-
ta: A. cand. CARDARIA draba: A. cand., P. paras. CARDUUS acanthoides,
crispus, nutans, personata albicans: B. lact. CARLINA acaulis: B. lact.
CARTHAMUS tinctorius: Pl. halstedit. CAUCALIS platycarpos: Pl. wmbel.
CENTAUREA atropurpurea: A. trag.; cyanus, jacea: B. lact.; kotschyana:
A. trag.; mollis, nigrescens: B. lact.; orientalis: A. trag.; pannonica
pannonica, phrygia carpatica, phrygia melanocalathia, phrygia phrygia,
phrygia pseudophrygia, rocheliana: B. lact.; ? scabiosa: A. trag.; sol-
stitialis, stenolepis, triumfetti: B. Zact. CENTAURIUM erythraea ery-
-thraea, pulchellum: P. erythraeae. CEPHALARIA laevigata: P. cephalartae-
554
laevtgatae; transylvanica: P. knauttae; uralensis multifida: P. cephala-
rtae—Laevtgatae. CEPHALORRHYNCHUS tuberosus: B. lact. CERASTIUM arvense,
brachypetalum: P. tomentosa; dubium, fontanum triviale: P. conferta;
glomeratum, semidecandrum: P. tomentosa. CERATOCARPUS arenarius: A. eu-
rottae, P. fartnosa. CERINHTE minor: P. certnthe. CHAENORRHINUM minus:
P. grtsea. CHAMOMILLA recutita: P. leptosperma; suaveolens: A. trag., P.
leptosperma. CHEIRANTHUS cheiri: A. cand., P. paras. CHENOPODIUM album,
ambrosioides, bonus-henricus, ficifolium: P. fartnosa; glaucum: P. che-
nopodtt-glauct; hybridum, murale, opulifolium, polyspermum, urbicum,
vulvaria: P. fartnosa. CHORISPORA tenella: A. cand., P. paras. CHRYSO-
SPLENIUM alternifolium: P. chrysosplentz.CICERBITA alpina: B. lact. CIR-
SIUM alatum: A. trag.; arvense, canum: A. trag., B. lact.; erisithales:
A. trag.; oleraceum: A. ‘trag., B. ltact.; palustre: B. lact. rivulare;
vulgare: A. trag., B. lact. CLEOME spinosa: A. cand., P. paras. COCHLE-
ARIA pyrenaica var. borzeana: P. paras. CONIUM maculatum: P. wnbel.
CONRINGIA orientalis: A. cand., P. paras. CONYZA canadensis: Ba. ento-
spora. CORISPERMUM nitidum: P. vtstulensts. CORONILLA scorpioides, va-
ria: P. corontllae. CORONOPUS squamatus: A. cand., P. leptdit. CORIDA-
LIS bulbosa marschalliana: P. corydalts; capnoides: P. corydalts-tnter-
medtae; solida: P. corydalts. CREPIS biennis, foetida rhoeadifolia, pa-
ludosa, pulchra: B. 2act.; sancta: A. trag., B. tact. ; setose: Baste.
CRUCIATA pedemontana: P. calotheca. CRUPINA vulgaris: A. trag. CUCUMIS
melo, sativus: Ps. cubensts. CYNOGLOSSUM hungaricum: P. cynoglosst.
DAUCUS carota: PZ. wmbel. DESCURAINIA sophia: A. cand., P. paras. DIGI-
TALIS grandiflora: P. grtsea. DIMORPHOTHECA pluvialis: B. lact.; sinua-
ta: Pl. halstedtt. DIPLOTAXIS muralis: A. cand., P. paras.; tenuifolia:
P. paras. DRABA lasiocarpa: A. cand.; nemorosa: P. paras. DIPSACUS ful-
lonum, laciniatus, sativus: P. dtpsact. ECHIUM vulgare: P. echtt. EPI-
LOBIUM hirsutum, palustre, parviflorum, roseum, tetragonum: Pl. eptlo-
bit. ERODIUM ciconium, cicutarium, hoefftianum: P. conglomerata. ERO-
PHILA verna s.1.: P. paras.; verna verna: A. cand. ERUCASTRUM nasturtii-
folia: P. paras. ERYSIMUM cheiranthoides: A. cand., P. paras.; comatum,
crepidifolium: A. cand.; cuspidatum, diffusum, odoratum, repandum, wit-
mannii: A. cand., P. paras. EUCLIDIUM syriacum: A. cand., P. paras. EU-
PHORBIA amygdaloides, cyparissias, esula tommasiniana: P. cypartsstae;
seguierana: P. valestaca; ? serrulata: P. euphorbtae. EUPHRASIA pecti-
nata: P. grtsea; rostkoviana, stricta: Pl. densa. FAGOPYRUM esculentum:
P. ducomettt.FERULAGO campestris: Pl. wmbel. FILAGO vulgaris: A. trag.
FOENICULUM vulgare: Pl. wnbel. FRAGARIA vesca: P. sparsa. FUMARIA of fi-
cinalis, schleicheri, vaillantii: P. affints. GALEGA officinalis: P. vt-
etae. GALIUM album, aparine, boreale, divaricatum, humifusum, mollugo,
odoratum, palustre, rubioides, schultesii, spurium, tricornutum, verum
verum: P. calotheca. GAZANIA rigens: B. lact. GERANIUM columbinum, dis-
sectum, lucidum, molle, phaeum: P. conglomerata; pratense: Pl. pustilq;
pusillum, robertianum: P. conglomerata; sylvaticum: P. conglomerata, Pl.
pustlla. GEUM aleppicum, coccineum, rivale, urbanum: P. sparsa. GLAUCI-
UM corniculatum: P. corydalts. GLYCINE max: P. manshurtca. GYPSOPHILA
muralis: P. gypsophtlae. HALIMIONE verrucifera: P. fartnosa. HELIANTHE-
MUM canum canum, nummularium s.1., nummularium nummularium, nummularium
obscurum , Oleandicum alpestre, salicifolium: P. leptoelada. HELIANTHUS
annuus : Pl. halstedtt. HELICHRYSUM bracteatum: B. lact. HELLEBORUS
Odorus odorus: Pl. pygmaea; purpurascens: P. ficartae, Pl. pygmaea. HES-
PERIS matronalis: A. cand., P. paras.; sylvestris velenovskyi: P. paras.
HIERACIUM argillaceum, aurantiacum, bifidum, caesium, caespitosum, cymo-
sum, ? lactucella, lactucella lactucella, murorum, pilosella micradeni-
um, pilosella pilosella, piloselloides piloselloides, praealtum s.1.,
Praealtum bauhinii, praealtum thaumasium, rotundatum, umbellatum: B.
Soo
Lact. HOLOSTEUM umbellatum s.1., umbellatum glutinosum: P. holostet.
HORNUNGIA petraea: A. cand. HUMULUS lupulus: Ps. hwnult. HUTCHINSIA al-
pina brevicaulis: A. cand. HYMENOLOBUS procumbens: A. cand. HYOSCYAMUS
niger: P. hyoscyamt. HYPOCHOERIS radicata: B. lact. IBERIS saxatilis:
A. cand.; umbellata: A. cand., P. paras. \IMPATIENS noli-tangere: Pl.
obducens. INULA britannica, conyza, salicina: A. trag. ISATIS tinctoria:
A. cand., P. paras. |ISOPYRUM thalictroides: P. parvula, Pl. pygmaea.
KNAUTIA arvensis: P. knauttae, P. vtclacea; ? dipsacifolia, drymeia dry-
meia: P. knauttae. KOCHIA scoparia: P. fartnosa. LACTUCA saligna, sati-
va, serriola, viminea: B. Lact. LAMIUM album, amplexicaule, garganicum
laevigatum, maculatum, purpureum: P. Zamtt. LAPPULA barbata, marginata,
squarrosa: P. echtnospermt. LAPSANA communis, intermedia: B. Lact. LA-
SERPITIUM latifolium: PZ. wnbel. LATHYRUS cicera, hirsutus: P. vtetae;
niger: P. sennentana; nissolia: P. vtctae; palustris: P. sennentana;
sphaericus, sylvestris: P. vtetae; tuberosus: P. sennentana. LEONTODON
autumnalis, hispidus s.].: B. lact. LEONURUS marrubiastrum: P. Zamtt.
LEPIDIUM campestre: A. cand., P. paras.; cartilagineum crassifolium: A.
cand ss densiflorum: A. cand., P. lepidit, P. paras.; latifolium, perfo-
liatum: A. cand., P. paras.; ruderale: A. cand., P. leptdtt. LEUCANTHE-
MUM nipponicum: P. leptosperma. LEVISTICUM officinale: Pl. wnbel. LI-
GUSTICUM mutellina: PZ. wnbel. LIMONIUM gmelinii, vulgare serotinum: P.
stattcts. LINARIA genistifolia genistifolia, vulgaris: P. grtsea, P. me-
Lampyrt. LINUM catharticum: P. Ztnz¢. LOBULARIA maritima: P. paras. LOG-
FIA arvensis: A. trag. LOTUS tenuis: P. Lotorwn. LUNARIA annua pachyr-
rhiza: A. cand., P. paras.; rediviva: P. paras. LYCHNIS alpina, visca-
ria: P. lyechnttts. MATRICARIA perforata: P. leptosperma, P. radtt. MAT-
THIOLA incana incana, longipetala bicornis: P. paras. MEDICAGO falcata:
P. aesttvalts; lupulina, minima: P. medtcagtnts-mintmae; sativa: P. ae-
sttvalts. MELAMPYRUM arvense, cristatum, saxosum: P. melampyrt. MELILO-
TUS alba, officinalis: P. aesttvalts. METHA longifolia: P. sttgmaticola.
MEUM athamanticum: PZ. wnbel. MOEHRINGIA trinervia: P. arenartae. MYCE-
LIS muralis: B. Lact. MYOSOTIS alpestris, arvensis arvensis, discolor
discolor, ramosissima ramosissima, scorpioides, sparsiflora, stricta,
Sylvatica sylvatica: P. myosottdts. NASTURTIUM officinale: P. paras.
NESLIA paniculata s.1l.: A. cand., P. paras. NICOTIANA alata, ipomopsi-
folia, langsdorfii, longiflora, nodiflora, paniculata, suaveolens, taba-
cum: P. tabacitna. QODONTITES verna s.1.: Pl. densa. OENANTHE silaifolia:
Bl. baudystt. ONOPORDON acanthium, tauricum: B. Zact. PAPAVER dubium: P.
arborescens; hybridum: P. ertstata; rhoeas, somniferum: P. arborescens.
PASTINACA sativa: PZ. wnbel. PELTARIA alliacea: P. paras. PETROSELINUM
crispum: PZ. wnbel. PEUCEDANUM alsaticum, austriacum, palustre: Pl. um-
bel. PHYTEUMA vagneri: P. phytewmatts. PICRIS echioides, hieracioides s.
1.: B. laet. PIMPINELLA major, saxifraga: PZ. umbel. PISUM sativum sati-
vum: P. vtetae. PLANTAGO altissima: P. alta; arenaria: P. plantagtntis;
cornutl, wanceolata: P, alta; major: P..alta, P.. plantagints; major in-
termedia, media s.1l.: P. alta. POLYGONUM arenarium arenarium, aviculare:
P. polygont. PORTULACA oleracea oleracea, oleracea sativa: A. portula-
cae. POTENTILLA anserina: PB. sparsa: aurea aurea, aurea chrysocraspeda:
P. potentillae; reptans: P. sparsa. PRIMULA veris: P. androsaces. RANUN-
CULUS acris acris, acris strigulosus, bulbosus, cassubicus, carpaticus,
constantinopoiitanus, crenatus, ficaria, flammula: P. f¢cartae; illyri-
cus: P. tllyrtca; lateriflorus, oreophilus, oxyspermus, pedatus, plata-
nifolius, polyanthemos polyanthemos, pseudomontanus, repens, sardous:
P. ftcartae. RAPHANUS raphaniastrum raphaniastrum: A. cand., P. paras. ;
raphaniastrum landra: P. paras.; sativus: A. cand., P. paras. RAPISTRUM
perenne: P. paras. RESEDA lutea, phyteuma: P. ertspula. RHINANTHUS alpi-
nus, angustifolius, minor: PZ. densa; rumelicus: P. melampyrt, Pl. den-
556
sa; wagneri: Pl. densa. ROCHELIA disperma retorta: P. rocheltae. RORIPPA
amphibia: A. cand., P. paras.; x armoracioides: P. paras.; austriaca,
islandica: A. cand., P. paras.; pyrenaica: A. cand.; sylvestris sylves-
tris: \A. cand., P. paras.; sylvestris kerneri: A. cand: ROSA gallicarar.
sparsa. RUBUS caesius, canescens, hirtus, idaeus: P. sparsa. RUMEX ace-
tosa, acetosella, alpestris, tuberosus tuberosus: P. rwntets. SALSOLA
kali s.l., kali ruthenica: P. vtstulensts. SANGUISORBA minor: P. sparsa.
SANICULA europaea: PZ. wnbel. SCABIOSA columbaria columbaria: P. knautt-
ae; columbaria s.1l.: P. vtolacea; lucida lucida, ochroleuca: P. knauttae.
SCLERANTHUS annuus, uncinatus: P. selerantht. SCORZONERA cana, hispanica,
laciniata: A. trag. SCROPHULARIA nodosa, scopoli: P. sordida. SENECIO
aquaticus barbareifolius, squalidus, subalpinus: B. lact.; vernalis: A.
trag., B. lact.; vulgaris: B. lact. SERRATULA tinctoria: A. trag. SETA-
RIA italica, verticillata, viridis: S. gramtntcola. SHERARDIA arvensis:
P. sherardtae. SILAUM silaus: PZ. wnbel. SILENE alba, noctiflora, oti-
tes: P. melandrit. SINAPIS alba, arvensis: A. cand., P. paras. S|ISYMBRI-
UM altissimum, loeselii, officinale, orientale, polymorphum: A. cand.,
P. paras.; strictissimum: P. paras. SIUM latifolium: Bl. baudystt. SMYR-
NIUM perfoliatum: PZ. umbel. SONCHUS arvensis s.1., arvensis uliginosus,
asper, asper glaucescens, oleraceus, palustris: B. lact. SPERGULA arven-
sis: P. obovata. SPERGULARIA marina, media: A. caryophytlacearum, P.
obovata; rubra: P. obovata. SPINACIA oleracea: P. fartnosa. STACHYS pa-
lustris: P. lamtt. STELLARIA holostea: P. parva; media s.1.: alstnearum.
SYMPHYTUM cordatum, nodosum, officinale s.l., officinale uliginosum, ot-
tomanum: P. symphytt. SYRENIA cana: A. cand.; montana: A. cand., P. pa-
ras. TANACETUM vulgare: P. leptosperma. TARAXACUM ‘officinale': B. Lact.
TETRAGONOLOBUS maritimus: P. tetragonolobt. THLASPI arvense, perfolia-
tum: A. cand., P. paras. TRAGOPOGON dubius: A. trag., Pl. sphaerosperma;
porrifolius, pratensis s.l.: A. trag. TRIBULUS terrestris: P. trtbultna.
TRIFOLIUM alpestre, arvense, badium, campestre, dubium, echinatum, hybri-
dum s.]1., hybridum elegans, medium s.1., medium banaticum, montanum, pra-
tense, repens, spadiceum, striatum: P. trtfoltorum. URTICA dioica, kio-
viensis: Ps. urttcae; urens: P. debaryt. VALERIANA officinalis: P. vale-
rtanae. VALERIANELLA dentata, lasiocarpa, locusta, pumila, rimosa: P. va-
lertanellae. VERBASCUM banaticum, blattaria, densiflorum, glabratum gla-
bratum, nigrum s.1., phlomoides, speciosum, thapsus s.1]., thapsus cras-
sifolium: P. sorditda. VERONICA anagallis-aquatica, anagalloides: P. gri-
S@a; arvensis, austriaca teucrium: P. sordtda; beccabunga, catenata: P.
grtsea; chamaedrys s.1l., hederifolia: P. sordida; officinalis: P. grtsea;
persica, polita, praecox, prostrata: P. sordtda; scardica, scutellata,
serpyllifolia serpyllifolia, serpyllifolia humifusa, spicata spicata,
urticifolia: P. grtsea. VICIA angustifolia, cracca, grandiflora: P. wi-
etae; hirsuta: P. ervt; lathyroides, lutea lutea, narbonensis, pannonica
S.]., pannonica striata, peregrina, pisiformis, sativa, sepium, tetra-
sperma, villosa: P. vtetae. VIOLA arvensis, kitaibeliana, tricolor tri-
color: P. vtolae. VITIS vinifera sylvestris, vinifera vinifera, vinifera
x American species: Pl. vtttcola. XANTHIUM ? italicum, strumarium: Pl.
halstedtt.XERANTHEMUM annuum: A. trag., B. lact.; cylindraceum: A. trag.
ZIZIPHORA capitata: P. lamit.
MYCOTAXON
VOurPeerVIy No.(2,-pp. 557-564 January-March 1983
Sa ee eee
SARCINULELLA BANKSIAE GEN. ET SP. NOV., A COELOMYCETE
WITH A UNIQUE METHOD OF CONIDIAL DISPERSAL
B.C. OULLON
Commonwealth Mycological Institute, Kew, Surrey, UK
and
J.L. ALCORN
Plant Pathology Branch, Department of Primary Industries, Meters Rd.,
Indoorooptlly, Queensland, Australia
SUMMARY
Saretnulella bankstae gen. et sp. nov. is described and
illustrated from bark of Banksta tntegrtfolta collected in
Queensland. The genus is characterized by pycnidial conidiomata
and phialidic hyaline aseptate conidia. Tendrils issuing from
the ostiole are comprised of catenate sacci of mucilage each
containing 40 or more conidia. The individual sacci retain their
integrity during dispersal. Periphyses lining the base of the
ostiolar channel are thought to be involved in saccus formation.
Amongst many interesting collections of microfungi made in
Queensland following the XIII International Botanical Congress
in Sydney, Australia, 1981, was a minute coelomycete. This
proved to have a conidial dispersal method previously unknown
in the Deuteromycotina.
Saretnulella gen. nov.
(etym. saretnula (L) - a little pack or bundle + ella (L)
diminutive)
558
Mycelium immersum. Conidiomata conica, separata, superficialia
vel semi-immersa, atro brunnea vel nigra; parietes crassi, ex
textura angulari brunnea sed in regione ostioli ex textura
porrecta verticali compositi; ostiolum centrale, circulare,
canali cylindrico ad basim periphysibus instructi. Conidiophora
absentia. Cellulae conidiogenae discretae, determinatae,
hyalinae, cylindricae, canalibus apicalibus phialidicis
conspicuis, spissescentibus periclinalibus distinctis sed collis
notatis absentibus , ex parietibus basalibus et lateralibus
conidiomatorum formatae. Conidia phialidica, hyalina, aseptata,
laevia, (1)-2 guttulata, ellipsoidea; plus 40 conidia in saccis
cylindricis aggregata, in catenis longis nonramosis e ostiolo
emergentia.
Sp. typ. S. bankstae
Mycelium immersed. Conidiomata conical, separate, superficial to
semi-immersed, dark brown to black; walls thick, of brown textura
angularis except in the ostiolar region which is of vertically-
orientated textura porrecta; ostiole central, circular, with a
cylindrical channel, the base of which is lined with periphyses.
Conidiophores absent. Conidiogenous cells discrete, determinate,
hyaline, cylindrical, each with an apical phialidic aperture,
channel, and periclinal thickening but without a collarette,
formed from the basal and lateral walls of the conidiomata.
Conidia phialidic, hyaline, aseptate, smooth (1)-2 guttulate,
ellipsoid; 40 or more grouped in cylindrical sacci which emerge
from the ostiole in long unbranched chains.
Saretnulella bankstaeé sp. nov.
Mycelium immersum. Conidiomata conica vel fere rostrata,
separata, superficialia vel semi-immersa, atro brunnea vel nigra,
laevia, nitida, usque ad 90 um diam x 150 um alta; parietes 15 um
crassi, ad basim stratum exteriorius ex textura angulari medio
brunnea, 4-5 cellulis crassis et stratum interiorius ex textura
angulari pallide brunnea, usque ad 10 cellulis parvioribus
crassis, faciebus ex textura angulari fuscioribus; canalis
ostioli cylindricus, 7-8 um diam, textura porrecta verticaliter
disposita brunnea circumcinctus, apicem versus pallidiore hyphis
plus separatis; periphyses hyalinae, deminutae, aseptatae, 10-12
x 1 um in fimbria ad basim canalis ostioli in lumen conidiomatis
projectae; ostiolum singulum, circulare, centrale, apicibus
libris hypharum texturae porrectae fimbriatum, 7-10 um diam.
Conidiophora absentia. Cellulae conidiogenae discretae,
determinatae, hyalinae, cylindricae, raro lageniformes, 4.5-7 x
2-3 um, canalibus apicalibus phialidicis conspicuis,
559
Fig. 1. Saretnulella banksiae. A, vertical median section of a
conidioma; B, conidia; C, conidiogenous cells lining the inner
face of the conidiomatal wall; D, ostiolar region of conidioma
with emerging sacci; E, vertical section of ostiolar region;
F, chain of sacci. D-F, conidia in sacci depicted at varying
optical levels.
560
Spissescentibus periclinalibus distinctis sed collis notatis
absentibus, ex parietibus basalibus et lateralibus conidiomatorum
formatae. Conidia 4-6 x 2-2.5 um, phialidica, hyalina, aseptata,
laevia, saepe 2-guttulata, raro l-guttulata, ellipsoidea; plus 40
conidia in saccis cylindricis, 14-21 x 8.5-9 um aggregatis, usque
ad 20 in catenis longis non ramosis e ostiolo emergentia.
In corticis emortuis Bankstae tntegrifoltae, Peregian Beach,
Queensland, Australia, B.C. Sutton & J.L. Alcorn, 29 Aug. 1981,
holotypus IMI 263392b; isotypus BRIP 13683.
Mycelium immersed. Conidiomata conical to almost rostrate,
separate, superficial to semi-immersed, dark brown to black,
smooth, shining, up to 90 um wide x 150 um high; walls several
cells thick, 15 um wide, at the base composed of an outer layer
4-5 cells thick of medium brown textura angularis and an inner
layer up to 10 cells thick of pale brown smaller-celled textura
angularis, at the sides of darker textura angularis; the ostiolar
channel is cylindrical, 7-8 um diam, and bounded by vertically
orientated brown textura porrecta which becomes paler, with the
hyphae more separated, towards the apex; at the base of the
ostiolar channel a fringe of hyaline, tapered, aseptate periphyses,
10-12 x 1 um, project into the lumen; ostiole 7-10 um diam, single,
circular, fringed by the free apices of the hyphae forming the
textura porrecta. Conidiophores..absent. Conidiogenous cells
discrete, determinate, hyaline, cylindrical, more rarely
lageniform, 4.5-7 x 2-3 um, each with an apical phialidic aperture,
channel, and periclinal thickening but no collarette, formed from
the basal and lateral walls of the conidiomata. Conidia 4-6 x
2-2.5 um, phialidic, hyaline, aseptate, smooth, often 2-guttulate,
occasionally l-guttulate, ellipsoid; 40 or more conidia are grouped
into cylindrical sacci, 14-21 x 8.5-9 um, which emerge from the
ostiole in long, unbranched chains comprising up to 20 individual
sacca.
DISCUSSION
In conidial morphology, conidiogenesis and some aspects of
conidiomatal structure Sareinulella differs little from many of
the known genera of coelomycetes (Sutton, 1980). Conidia are
comparatively small, unicellular and hyaline; they are formed
from short cylindrical conidiogenous cells which show a typical
apical phialidic apparatus. The conidiomata are pycnidial with
a single locule and walls are several cells thick. It is in the
organization of the ostiolar tissues and the method of dispersal
of conidia that Saretnulella is quite unusual.
Comparatively few genera of coelomycetes have rostrate conidiomata
in which the walls surrounding the ostiolar channel are of textura
561
Fig. 2. Saretnulella bankstae. A,C, chains of sacci containing
conidia (A, x 1000; C, x 650); B, ostiolar region of conidioma
with emerging sacci ( x 650).
502
porrecta, (Sutton, 1973, 1975, 1980). It is only in Nanoschema Sutton
and Cornucoptella Hohn. where loose textura porrecta forms the upper
part of the conidioma and even in these genera the ostiole and ostiolar
channel are not well-differentiated. In Saretnulella the channel is up
to 65 um long and the tissue surrounding it becomes progressively less
compact towards the apex though still retaining its structural
integrity. This results in an ostiolar channel of constant diameter
throughout its length. At the base of the channel a fringe of short
sterile periphyses projects into the lumen. Although paraphyses
interspersed with conidiophores are documented for a few genera of
coelomycetes (Sutton, 1980 ; Nag Raj, 1981), periphyses are even less
frequent. Nag Raj & DiCosmo (1978) report the ostiole in Melancontopsts
tnquinans Ell. & Ev. as lined with simple or branched, septate,
subhyaline, smooth-walled, sterile hyphae invested in a mucilaginous
matrix. Apart from this example Saretnulella appears to be the only
other coelomycete in which such sterile elements have been confirmed.
The mechanism of conidial dispersal in Saretnulella is quite remarkable.
The majority of coelomycetes produce conidia in slimy mucilaginous
material, and under suitable conditions of temperature and especially
humidity, the conidia suspended in the mucilage are dispersed through
the ostiole, raphe, irregular tear or overlying host tissues in either
distinct tendrils (as in Cytospora Ehrenb.: Fr. or Ltbertella Desm.) or
in effuse or compact conidial masses (as in Phoma Westd. and Me Laneontum
Lk: Fr.). The common features of these different modes of dispersal is
that the conidia lie at random and are not particularly orientated in
any direction in the supporting mucilaginous material. On mounting
conidial tendrils or masses in aqueous or phenolic preparations, the
mucilage dissipates, thus allowing the conidia to become scattered. In
Saretnulella however, the conidia fill the conidiomatal locule and
subsequently become aggregated into groups of 40 or more in the ostiolar
channel after passing the periphyses. The individual sacci of conidia
emerge from the ostiole in long unbranched chains which, even under the
dissecting microscope, can be seen as quite different from normal
conidial tendrils because of their segmented appearance. When chains
are prepared in aqueous or phenolic mountants they may break at the
junctions between sacci but the individual sacci do not lose their
integrity. Erythrosin, acid fuchsin, trypan blue and cotton blue fail
to stain either the mucilaginous sacci or the contained conidia.
It seems most probable that the periphyses are involved either in
production of mucilage or the chemical modification of mucilage formed
in the conidiomatal locule because sections of conidiomata always show
the locule to be filled to capacity with conidia, and these are randomly
distributed. It is only after passing the lining of periphyses that a)
the conidia become aggregated into sacci and b) the mucilage becomes
altered to form the rigid cylindrical matrix in which the conidia are
held, The diameter of the ostiolar channel clearly imposes a structural
limitation on the width of the conidial sacci, but the mechanism(s)
regulating the length of sacci is not known. Conidial sacci vary from 14
to 21 um in length and a combination of temperature, humidity and diurnal
periodicity may be involved in determining not only the numbers of
conidia produced but also the length of the sacci.
563
Reports of the aggregation of conidia in this very precise manner
have not so far been made for any genera of coelomycetes or
hyphomycetes other than Saretnulella. To differentiate this type of
arrangement from that known in the majority of slimy-spored
coelomycetes and many hyphomycetes, and from production of strong
and weak chains of connected and disconnected conidia (Minter et al.,
1982, 1983), it is proposed to introduce the term saccus (sac, L).
This describes the individual cylindrical packets of conidia bound in
rigid mucilaginous material. They are distinct from the random
suspension of conidia in mucilaginous tendrils or effuse masses formed
by other fungi, and which vary according to the humidity to which the
fungus may have been subjected.
The sacci in Sarcinulella raise interesting questions concerning
production of conidia, their dispersal and germination, and the
establishment of new mycelium capable of forming conidiomata. What is
the composition of the mucilaginous matrix in which conidia are
dispersed and how does it differ from that in other coelomycetes and
hyphomycetes? How and where is it formed? What factors determine the
length of the sacci and the number of conidia included in each saccus?
Initial formation of conidia from phialides is accompanied by production
of normal mucilage but the elements for dispersal, the sacci, may well
be dry and are presumably dispersed by wind. It is possible that the
sacci may be moist and therefore sticky if humidity is high. Do these
factors give an increased survival value to this method of dispersal,
and if so why is it apparently unique? Once the sacci have been
dispersed how is the matrix broken down and what proportion of conidia
are capable of germination? Such questions cannot be effectively
answered unless more material is obtained and the organism is brought
into culture and stimulated to sporulate. Further efforts in these
directions are being made,
B.C. Sutton is grateful to the Department of Primary Industries,
Queensland and the Rural Credits Development Fund for financial support
during his stay in Australia in 1981. Both authors thank Miss C. Osborn
and Mr. D. Fry for their technical and photographic expertise
respectively.
REFERENCES
Minter, D.W., Kirk, P.M. & Sutton, B.C. (1982). Holoblastic phialides.
Transactions of the British Mycological Society 79 : 75-93.
Minter, D.W., Kirk, P.M. & Sutton, B.C. (1983). Thallic phialides.
Transactions of the British Mycological Society 80 (in press).
564
Nag Raj, T.R. (1981). Coelomycete systematics. In Biology of Conidial
Fungi 1 (ed. G.T. Cole & B. Kendrick). pp. 43-84. Academic
Press.
Nag Raj, T.R. & DiCosmo, F. (1976). Icones generum coelomycetum fasc.
X. University of Waterloo Biology Series 19 : 1-45.
Sutton, B.C. (1973). Coelomycetes. In The Fungi IVA (ed. G.C.
Ainsworth, F.K. Sparrow & A.S. Sussman), pp. 513-582.
Academic Press.
Sutton, B.C. (1975). Eucalyptus microfungi. Satchmopsis gen. nov. and
new species of Coniella, Coniothyrium and Harknessia. Nova
Hedwigia 26 : 1-16.
Sutton, B.C. (1980). The Coelomycetes, pp. 1-696. CMI, Kew.
MYCOTAXON
VOI EAVA, Now. 2 Dp. SOS—571 January-March 1983
CULTURAL STUDIES IN HYPOCHNICIUM
(CORTICIACEAE, BASIDIOMYCETES)
NILS HALLENBERG
Department of Plant Taxonomy
Universtty of Gdteborg, S-413 19 Géteborg, Sweden
ABSTRACT Hypochnicium karstenii (Bres.) Hal lenb.. nov.
comb. is segregated from H. bombycinum (Sommerf. ex Fr.)
J@jeerikss: on the basis of incompatibility.
inher closely related species H. eichleri (Bres.)) Erakse.
e Ryv'.. and HH.) punctulLatum. (Cke). John Erikss. anevtound
to be incompatible.
Hypochnicium bombycinum (Sommerf. ex Fr.) John Erikss.
has been treated as one taxon with a noticeable internal
variation (Erikss. & Ryv., 1976): The hymenium may be
smooth to more or less tuberculate or even irpicoid and
the spores may be characteristically ovoid or ellipsoid -
- subglobose. In. N. Europe, this variation in morphology
seems to be correlated to the occurrence on different
kinds of substrates.
Specimens collected on Pinus or Picea have membranaceous
fruitbodies and a high ratio of ellipsoid-subglobose
spores compared with the ovoid spore shape. Such specimens
have been described as H. bombycinum var. pinicola Lundell
(Lundell & Nannfeldt, 1941).
The collections from deciduous wood are usually more crus-
taceous and varyingly tuberculate. Spores with an ovoid
shape are almost as common as ellipsoid-subalobose ones.
No other differences in basidiocarp morphology have been
found. For further descriptions of these taxa, see Eriks-
son and Ryvarden (1976).
In the present study it is shown that var. pinicola is
incompatible with var. bombycinum and thus represents an
independent species. Hypochnicium bombycinum var. pini-
cola has earlier incorrectly been referred to Corticium
molle Fr. (Karsten, 1882) - a species nowadays placed in
Leucogyrophana. Bresadola (1911) noticed this missappli-
cation and described Corticium karstenii Bres. as a new
species.
566
Hypochnicium karstenii (Bres.) Hallenb. n. comb. is here-
with proposed as a new combination. Basionym: Corticium
karstenmir ores... Ann. myc, 92425. C19 A Ee
Holotype: Mustiala, in corticem Pini, Karsten (S).
Holotype of H. bombycinum (Sommerf. ex Fr.) John Erikss.
is the following specimen: Thelephora bombycina, Salt-
dalen ("Saltadalen"), in cortice subpute. Alni dics.
Sommerfelt (0).
A closer investigation of the variation in spore size
and shape has been made (Fig. 1).
inchs skarstenaga’ Sehnetspores: are, J=——10x6<4 5-3 om,
average 8.7x7.6 pm; in H. bombycinum 7.5—11x6—7.5 pm,
average 8.7x6.3 pn.
In H. karstenii, the ratio ellipsoid/ovoid spores varies
between 2,0—4'3* in? H. ‘bombycinum 1) =) 4535:.
In the tests for extracellular oxidases, H. karstenii
gave negative results, thus differing clearly from H.
bombycinum. Concerning other cultural characteristics, no
noticeable differences could be found.
QD
0 OO
Fig. 1. A) Ellipsoid - subglobose spores in Hypochnicium
karstenii GB 79. B) Ovoid spores in H. bombycinum. GB 253..
567
MATERIALS AND METHODS. Monosporous mycelia have been ob-
tained from the following specimens:
Guiktaiee wo.
GB 37 Hypochnicium bombycinum/Sweden, Vastergétland,
G6teborg Botanical Garden/1974—08—15/ T. Halling-
back.
GB 253 H. bombycinum/Sweden, Torne Lappmark, between
Bjorkliden and Abisko/On\a leaning log iof /)Salix/
1981-08—11/ N. Hallenberg 3766.
GB 66 He karsteni1/Dennark, Jutland, Mols, Bierge, iN. -of
Tinghulen/On a leaning log of Pinus/1979—10—16/
VN... Hallenbers 315.3:
GB 79 H. karstenii/Sweden, Ostergétland, Omberg, Stor-
pissan nature reserve/1979—11—09/ K. Hjortstam
ieiesOblne
Two monosporous mycelia from each specimen were crossed
in all combinations. The mycelia were checked for the
presence of clamps since they had been in contact with
each other for 4 weeks.
GB 37/1 GB 37/2 GB 66/1 GB 66/2 GB 79/1 GB 79/2 GB 253/1 GB 253/2
GB
GB
GB
GB
GB
GB
GB
GB
CULTURAL CHARACTERISTICS FOR H.
37/1
37/2
66/1
66/2
79/1
79/2
253/1
253/2
+
+
+
+
-
+
BOMBYCINUM (GB 253).
Margin submerged, mat appressed, aerial mycelium absent;
hyphae with clamps, ordinarily branched, thin-walled,
2—5 pm in diam.
NOBLES' CODE (Nobles 1965, with emendations by Boidin,
13 6.60v8
C2 ee OC odes
2a.
36.
385 45.554.
>e.
6.1
568
OXIDASE REACTIONS:
gallie “acid + tyrosine -
guaiacol +++ para-cresol L (milk white)
Gradation accarcding "to Boidin C1958).
The number of monosporous mycelia present in 37 and 253
was too small to evaluate the polarity: Boidin (1956) re-
ports H. bombycinum var. trpicoitdes Bourd,. & Galz. as ter-
rapolar. Confrontations between the latter and GB 37, 66,
and 253 gave negative results. However, the still present
strain of H. bombycinum Var. “irpicoides’ (LY 1402/C) was
probably too old to allow any conclusions about the compa-
pte Rl om gl Ba bine
CYTOLOGY: Spores are uninucleate, monosporous mycelia ani-
nucleate, and polysporous mycelium regularly dicaryotic.
Staining methods according to Boidin (1958).
CULTURAL CHARACTERISTICS FOR .Hs ‘KARSTENII “(GB 66),
The investigated polysporous mycelium was made artificially
by inoculating the 6 available monosporous mycelia on the
same plate. Nodose-septate mycelium was isolated after 4
weeks.
Margin submerged, mat appressed, aerial mycelium sparsely
present, downy; hyphae with clamps, ordinarily branched,
thin-walled, 2-7 pm in diam.
NOBLES' CODE;
Li BC ee, OO. etree as. 1D OU cee Os lire
OXIDASE REACTIONS: No staining occurred in media containing
galilic acid, guaiacol, tyrosine or para=cresol.. However,
when drop-tests (according to Currie D. Marr, 1979) were
used, a very weak staining occurred with guaiacol and 1-
-naphtol (after 2 hours). More oxidase tests with true
polysporous mycelia ought to be done to make this result
more certain.
POLARITY: A,B: 1 A,B: 3/5780
A, Bo: 4 A5Bo: 2
CYTOLOGY: Spores are uninucleate, monosporous mycelia uni-
nucleate.
= OOO maa
Hypochnicium eichleri (Bres.) Erikss. & Ryv. is very close
to H. punctulatum (Cke) John Erikss. Compatibility tests
between the species have long been desired to ascertain
509
their independency from each other. Eriksson and Ryvarden
(1976) keep them apart on the following basis:
H. eichleri has a spore size of 8—10 (—12) x6—7 (-—7.5) pam;
the main distribution area in Europe is in the N. parts.
H. punctulatum has spores of a smaller size (5—)5.5-6.6
(—7.7)x4.5-5 wm. It is widely distributed in C. Europe
and with very few records in the N. parts.
Both species have been collected on coniferous as well as
deciduous wood. For further descriptions of the species,
see Eriksson and Ryvarden (1976).
In’ this study at is shown. that H. eichleri is incompatible
with H. punctulatum. H. punctulatum is found to be hetero-
thallic and tetrapolar, while H. eichleri seems to be ho-
mothallic. Further, the culture tests show that the myce-
lium in H. punctulatum grows faster than in H. eichleri.
Hyphal tips in H. eichleri are partly sulfo-positive,
while not stained at all in H. punctulatum.
MATERIAL AND METHODS. The following specimens were used
in culture tests:
Gulseuwce nor.
GB 29 Hypochnicium eichleri/Sweden, Vadstergétland,
Géteborg, Rya Skog nature reserve/1973—12-—11/
ge talline back.
GB 269 H. eichleri/Sweden, Torne Lappmark, Jieprenjakk/
On a fallen brown-rotted log of Betula/1981—08—
-09--10/ N. Hallenbere 3688.
GB 160 H. punctulatum/Denmark, Lolland, Fuglesang Stor-
skov/On a fallen log of Abies/1980—09—-18/ N.
Hallenberg 3406.
5 monosporous cultures from 269 were regularly clamped and
dicaryotic after 7 weeks, as well as the only one avail-
ape frome Ge (29). whis indicates. that H.,eichveri. might be
homothallic.
The identity of H. eichleri as a species distinct from
H. punctulatum is confirmed by di-mon tests between
clamped, dicaryotic cultures from H. eichleri’ and mono-
sporous, haploid cultures from H. punctulatum:
Dicaryotic x haploid cultures
Coeg29/ 1 °* GB-1600/2 = =
GEezZ69/1 x GB 160/1 = —
GB 26971 x GB 160/4 = -
GBE209/2 x GBR 160/1 .= -
Gee26 9/2 x GB 160/4: = =
In all confrontations clamps were completely absent in the
haploid side and a barrier zone was developed between the
mycelia.
CULTURAL CHARACTERISTICS FOR Hs /ELCHUERT. (GB 2690-2
Mat appressed, downy; hyphae with clamps at all septa, or-
dinarily branched, thin-walled, in the margin 4—-7.5 pm in
diam., in old parts of the mycelium 2—8 pm in diam., part-
ly widened to gloeocystidium-like structures, up to 50x8 pm.
After 6 weeks growth on an agar plate, some parts of the
hyphae were sulfo-positive — especially the hyphal tips.
CYTOLOGY: Spores uninucleate, monosporous mycelia dicaryo-
tic.
NOBLES' CODE:
2a. M2b) Ro Se. WIZ (26) eo 2h OL Do. 4 420.4 meen onteS
OXIDASE REACTIONS:
Catworrac dd +++ tyrosine -
guaiacol ++++ pave -cresco lL
CULTURAL CHARACTERISTICS FOR H. PUNCTULATUM (GB 160).
Margin submerged, mat appressed, aerial mycelium downy;
hyphae with clamps, 2—5 yum in diam., the wider hyphae
straight, sparsely septated, from them branching densely
ramified, narrow hyphae. In old parts of the mycelium some
hyphae are irregularly widened, forming gloeocystidium-
-like structures, 22—42x5-—8 pm. All parts of the mycelium
are sulfo-negative.
POLARAT YX: A,B: 1 A,Bo: 4
A,B): 5 A,B: 2
CYTOLOGY: Spores uninucleate, monosporous mycelia with
uninucleate cells, polysporous mycelia dicaryotic.
NOBLES' CODE:
2a 3G. cose oeteoO. 30. 45,0547 05.) OUNILOt,
OXIDASE REACTIONS:
ga bivice ae ad +++++ tyrosine ~
guaiacol +++ + para-cresol =
5 ae
DISCUSSION. Both Hypochnicium karstenii and H. punctulatum
are tetrapolar. This is in accordance with previous re-
sults in the genus, i.e. H. bombycinum var. irpicoides
Bourd. & Galz. (Boidin, 1956), H. cystidiatum Boid. 8
Gilles (Boidin and Lanquetin, 1971), and H. vellereum (E01
& Crag.) Parm. (Nobles and Nordin, 1955). On the contrary,
it is indicated that H. eichleri is homothallic. Hyphal
tips in cultivated mycelium of H. eichleri were found to
be sulfo-positive — a reaction previously found only in
the basidiocarps (gloeocystidia) of H. analogum (Bourd.
& Galz.) John Erikss. within the genus.
ACKNOWLEDGEMENTS
I am most grateful to Dr. Paule Lanquetin, Lyon, for valuable dis-
cussions. Gunnel Johansson, Géteborg, has kindly checked the English.
REFERENCES
Botdin td. 71956.) Polarité dite, "sexuelle wet "systématique
chez les Basidiomycétes Théléphoracés. Rev. Mycol.
2A C2=—3)e 1 24=1.3.1).
1958. Essai biotaxonomique sur les Hydnés re-
supines et les Corticiés. Rev. Mycol. Mem. hors,
Sevie6, 3838 opp.
1966. Basidiomycétes Corticiaceae de la Ré-
publique Centrafricaine. I, Le genre Gloeocystiellum
Donk. Gah. Maboké) 441) 2..5—174
Boidin.) J.) Lanqwetin; Paule. 19714. Basidiomycétes Corti=
ciaceae de la République Centrafricaine. III, Le
genre Hypochnicium Erikss. Cah. Maboké 9(2): 89-93.
Bresadola J..1911. Adnotanda mycologicas Ann. myc.) 9:
425-428.
ELSON, Js guRkyvarden, .L.4.1976.' The iCorticiac cae! of Worth
Burope.. Vol. .04. Sungat bor agen) Os lio,
Karsten. PA. 1882. Hattsvampar. 11. Bidr. Finl. nat folk
Bye ClO) 22 Si]. pp.
Pundeid ~ S.40 Nannfeldt, J.A. 1941. Fung. exs.csuecs Fasce
Rs Ow 1.052..
Marcuse Curie DD, 1979... Laccase and tyrosinase osidationsor
spot, test teagents. Mycotaxon TX (1).) 244-276).
Noptess.M-Ks 19654 Identification of cultures of wood-i1n-
habiting Hymenomycetes. Can. Journ. Bot. 43: 1097 —1139.
Nobles, M.K., Nordin, V.J. 1955: Studies in wood-inhabiting
Hymenomycetes II, Corticium vellereum Ell. & Crag.
Cane Uris 2b0G. 32) LO5—1.12.
MYCOTAXON
VOL OXxV UNG. eee hp aoe January-March 1983
NEOs Care
MSA PLANS FOR IMC
The Mycological Society of America has announced their
appointment of Garber Travel, Box 404, Brookline; iin
02146 as their official agents for travel arrangement to
the Third International Mycological” Congress to) bemmierd
in Tokyo, Japan, August 28-September 3, 1983. A brochure
is available detailing a Congress Program (August 26-
September 4th) only, a Congress and Japan Pre-tour (Aug-
ust 20-September 4th), and a Congress and China Pre-
tour (August 18-September (4th). Connections for those
also planning on attending the Fourth International Con-
oress “on “Plant Pathology” in Melbourne, Australiana,
August. are also arranged by Garber Travel: Because
of airline. pricing, air fares for. the (\CongressSmirccgag
alone are higher ($1413) than for the Congress plus Japan
Pre-tour ($961) or the Congress plus China Pre-tour
($1196). For those who wish to stay in Japan beyond 14
days, Garber Travel will arrange an APEX air fare of
$930. (All prices based on West Coast departures.)
The Mycological Society of America will also be awarding
a limited number of Travel Grants, each in the amount
of $500, to qualified members of the Society attending
and participating in the Tokyo meetings: Particwiar.
concerning elegibility and dates of application will be
announced in mid-January, 1983, by Dr. R. L. Shatter
Chairman, MSA Travel Awards Committee for IMC, The
Herbarium, University of Michigan, Ann Arbor, MI 48109,
in a mailing to MSA members residing in the U.S. Should
MSA members not have received information on application
procedures by mid-February, they should write to Dr.
Shaffer directly.
Bi
AUTHOR INDEX, VOLUME SIXTEEN
ACSAT, JAN & DAVID L. LARGENT. Ectomycorrhizae of selected conifers
growing in sites which support dense growth of bracken fern 509-518
ACSAI, JAN & DAVID L. LARGENT. Mycorrhizae of Arbutus menztesti Pursh.
and Aretostaphylos manzantta Parry in northern California 519-536
AJELLO, LIBERO, see McGINNIS §& AJELLO
ALCORN, J.L. On the genera Cochliobolus and Pseudocochltiobolus 353-379
ALCORN, J.L., see PEGG § ALCORN, see SUTTON § ALCORN
ALFIERI, SALVATORE A., JR., see SCHOULTIES & al.
AWAO, TAKEYOSHI & SHUN-ICHI UDAGAWA. A new thermophilic species of My-
celtophthora 436-440
BAZZALO, MARTA E. & JORGE E. WRIGHT. Survey of the Argentine species of
the Ganoderma luctdum complex 293-325
BEN-ZE'EV, ISRAEL. Erynta neopyraltdarum sp. nov. and Contdtobolus apt-
culatus, pathogens of pyralid moths: components of the misdescribed
species Entomophthora pyralidarum [Zygomycetes, Entomophthorales] 273-
292
BINYAMINI, N. Tremellales of Israel 380-386
BOIDIN, J. & P. LANQUETIN. Basidiomycétes Aphyllophoralés epitheloides
étalés 461-499
BRAUN, UWE. Descriptions of new species and combinations in Microsphaera
and Erystphe (III) 417-424
BRAUN, UWE. Taxonomic notes on some powdery mildews (II) 425-428
CONSTANTINESCU, O. §& G. NEGREAN. Check-list of Romanian Peronosporales
537-556
CRANE, J.L. & ADRIANNA D. HEWINGS. Stilbellaceous fungi 1. Didymosttlbe
133-140
DAS, B.K., see YEN € al.
DISSING, HENRY §& SIGMUND SIVERTSEN. Operculate Discomycetes from Rana
(Norway) 5. Rhodoseypha gen. nov. and Rhodotarzetta gen. nov. 441-460
DUNN, MICHAEL T. A new species of Endophragmiella from sclerotia of
Selerotinta minor 152-156
EGGER, K.N., see HAINES §& EGGER
EL-GHOLL, NABIH E., see-SCHOULTIES & al.
GALLOWAY, D.J, see RENNER §& GALLOWAY
GLAWE, D.A., C.B. SKOTLAND §& W.J. MOLLER. Isolation and identification
of Eutypa armentacae from diseased grapevines in Washington State 123-
E32
GRATTO, KELLY A. & DARRYL W. GRUND. Paraffin embedding and semithin sec-
tioning of basidiocarp tissues 239-242
GRUND, DARRYL W., see GRATTO §& GRUND
GUZMAN, GASTON. New species of fungi from the Yucatan Peninsula 249-261
HAINES, J.H. §& K.N. EGGER. A new species of Orbilta from Canada 107-113
HALE, MASON E., JR. A new species of Parmelta (Lichenes) with protoce-
traric acid 162-164
HALE, MASON E., JR., see PIKE & HALE
HALLENBERG, NILS. Cultural studies in Hypochnicitum (Corticiaceae, Basi-
diomycetes) 565-571
HAWKSWORTH, D.L. & M.A. SHERWOOD. Two new families in the Ascomycotina
262-264
HENNEBERT, G.L. Revue des Livres 331-334
HEWINGS, ADRIANNA D., see CRANE §& HEWINGS
JENKINS, DAVID T. A new species of Amantta 414-416
KAR, A.K., see YEN @& al.
KAUSHAL, RISHI. A reinvestigation of the north-west Himalayan Pulvinulas
117-122
KOHN, L.M. A preliminary discomycete flora of Macaronesia: Part 5, Scle-
rotiniaceae 1-34
574
KORF, RICHARD P. Sanctioned epithets, sanctioned names, and cardinal
principles in; > Pers.” and "2 Pr.) Citations Gadeooe
KORF, RICHARD P., see SHARMA & KORF
KOSKE, R.E, DIANE DOUD MILLER §& CHRISTOPHER WALKER. Gtgaspora rettcula-
ta: a newly described endomycorrhizal fungus from New England 429-435
LANQUETIN, P., see BOIDIN §& LANQUETIN
LARGENT, DAVID L., see ACSAI §& LARGENT
LICHTWARDT, ROBERT W., see WILLIAMS @ al.
LIM, GLORIA, see YEN §& LIM
McGINNIS, MICHAEL R. & LIBERO AJELLO. A note on Sporotrichum gougerotit
Matrouchot 1910 232-238
MILLER, DIANE DOUD, see KOSKE & al.
MOLLER, W.J., see GLAWE & al.
MORGAN-JONES, G. Notes on hyphomycetes. XLII. New species of Acrodtctys
and Pseudosptropes from South Africa 187-191; XLIII. Concerning Chaeto-
pstna romanttca 192-196
MORGAN-JONES, G. & JAMES F. WHITE. Studies in the genus Phoma. I. Phoma
amerteana sp. nov. 403-413
NEGREAN, G., see CONSTANTINESCU §& NEGREAN
PARMASTO, ERAST & ILMI PARMASTO. Variability of spores of different ba-
Sidiocarps growing on the same mycelium in the Aphyllophorales 141-151
PARMASTO, ERAST & ILMI PARMASTO. Basidiospore variation in local popula-
tions of some Aphyllophorales 243-248
PARMASTO, ILMI, see PARMASTO §& PARMASTO
PEGG, K.G. & J.L. ALCORN. Phytophthora operculata sp. nov., a new marine
fungus 99-102
PENNYCOOK, S.R. Colletotrtechum gloeosportotdes (Penzig) Penzig et Sac-
cardo 507-508
PETERSON, STEPHEN W., see WILLIAMS @ al.
PFISTER, DONALD H. A new host for Sphaeronaemella helvellae — Pseudo-
rhiztna sphaerospora 165-166
PIKE, LAWRENCE H. §& MASON E. HALE, JR. Three new species of Hypogymnia
from western North America (Lichenes: Hypogyniaceae) 157-161
PLANK, S. & L. RYVARDEN. Phelltnus andina Plank & Ryv. nova sp. 114-116
RAJCHENBERG, MARIO. New South American resupinate polypores 500-506
RAJCHENBERG, M., seé@ RYVARDEN @ al.
RENNER, B. §& D.J. GALLOWAY. Phycosymbiodemes in Pseudocyphellarta in New
Zealand 197-231
RYVARDEN, L., J.E. WRIGHT & M. RAJCHENBERG. Megasporoporia, a new genus
of resupinate polypores 172-182
RYVARDEN, L., se@ PLANK §& RYVARDEN
SCHOULTIES, CALVIN L., NABIH E. EL-GHOLL §& SALVATORE A. ALFIERI, JR. Cy-
lindrocladitum spathtphyllt sp. nov. 265-272
SHARMA, R. & RICHARD P. KORF. Two new species of Helotiales from the
eastern Himalayas 326-330.
SHERWOOD, M.A., see HAWKSWORTH §& SHERWOOD
SIVERTSEN, SIGMUND, see DISSING § SIVERTSEN
SKOTLAND, C.B., see GLAWE & al.
STATES, JACK S. New records of hypogeous Ascomycetes in Arizona 396-402
SUTTON, B.C. & J.L. ALCORN. Sarcinulella bankstae gen. et sp. nov., a
Coelomycete with a unique method of conidial dispersal 557-564
UDAGAWA, SHUN-ICHI, see AWAO §& UDAGAWA, see UEDA §& UDAGAWA
UEDA, SEIICHI §& SHUN-ICHI UDAGAWA. A new Japanese species of Neocosmo-
spora from marine sludges 387-395
VANKY, KALMAN. Thecaphora androsacina and Entyloma gatllardtanum, new
species of Ustilaginales 103-106
WALKER, CHRISTOPHER, see KOSKE & al.
WHITE, JAMES F., see MORGAN-JONES §& WHITE
WILLIAMS, MARVIN C. Smttttum cellaspora, a new Harpellales (Trichomy-
cetes) from a chironomid hindgut 183-186
WILLIAMS, MARVIN C., ROBERT W. LICHTWARDT & STEPHEN W. PETERSON. Smtttt-
um Longtsporum, a new Harpellales (Trichomycetes) from chironomid guts
167-171
WRIGHT, JORGE E., see BAZZALO §& WRIGHT
WRIGHT, J.E., see RYVARDEN & al.
YEN, JO-MIN, A.K. KAR & B.K. DAS. Studies on hyphomycetes from West Ben-
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II. Cereospora and allied genera of West Bengal, 2 58-79; III. Cerco-
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376
INDEX TO FUNGOUS AND LICHEN TAXA, VOLUME SIXTEEN
This index includes genera, infrageneric taxa, species, and infra-
specific taxa. New taxa are in
boldface as are the pages where
they are published. New suprageneric taxa are in bold CAPITALS.
The fungus/host "Check-list of
Romanian Peronosporales" is itself
an index, reference to which is indicated by "see 540-556."
Acaulospora 435; bireticulata 435
Acremonium 393, 394; brunnescens
393; hyalinulum 393; tenuicristatum 387
Acrodictys 187, 189; appendiculata
189; brevicornuta 189; eickerii 187-189
Agaricus melleus 348, 349; obscurus
348
Albugo 539, see 540-556
Aleuria aphanodictyon 458
Amanita 414; sect Lepidella 414-416;
sect Validae 414-416; antillana 258; de-
missa 416; dunicola 249, 256, 257; ela-—
ta 255; gemmata 519; media 414-416;
polypyramis 416; praegraveolens 256;
Sukwulen: QS, P35, Poy, 255, MSs “Sic
4" 256; “sp. 5'' 258; yucatanensis 249,
(Sy PAS)
Amanitopsis floridana 258
Amylosporus 500, 501; campbelli 501;
wrightii 500, 501, 503
Antrodia 505; lindbladii 505
Anzia jamesii 204
Armillaria 348, 349; mellea 348
Armillariella 348; mellea 348
Arthrocristula hyphenata 155
Ascodichaena 262, 263
ASCODICHAENACEAE 262
Asterostroma 465
Auricularia auricula 385
Balsamia magnata 396, 397
Basidiophora 539, see 540-556
Bipolaris 353-355, 359-364, 367, 369-
373; australiensis 361; bicolor 355; chlo-
ridis 353, 373; coicis 367, 369; cynodon-
tis 355; dactyloctenii 355; hawaiiensis
355; shomomorphus) 353, 355,, 373,. 374;
indica 363, 367; kusanoi 355; maydis
255. 3593. 360,362, 363, 3673 melinidis
355; nodulosa 355; oryzae 355; peregi-
anensis 355; perotidis 355; ravenelii
355, 361; setariae 355; sorokiniana 355,
361-363, 367; spicifera 355, 361; steno-
spila 355; tripogonis 355; victoriae 355;
zeicola 355
Bisporella 326, 328; calycellinoides
326-328
Blumeria graminis 428
Boidinia 492
Boletus aureus 520, 522; dimidiatus
297; laccatus 297; lucidus 297; striatus
351
Botryotinia 6, 9
Botrytis 6, 9
Bremia 539, see 540-556
Bremiella 539, see 540-556
Bryodiscus 262, 263
Bryophagus 264
Calocera cornea 383, 385
Calocybe 520, 521
Calonectria kyotensis 271
Calostilbella calostilbe 139
Calycellina 328
Coenococcum graniforme 510, 512, 513,
So, SS, 520, S22, 2s
Cephalosporium 393; proliferatum 393,
394
Cercoseptoria caesalpiniae 80, Slee
cedrelae 80, 82, 83; cordiicola 96, 98;
vignicola 35, 37
Cercospora 35, 58, 72, 802 agawalia
70; alternantherae-nodiflorae 41; atri-
dis 60; bouganvilleae 98; coffeae 72;
coffeicola 72; cordae 98; cordiicola 98;
daturina 41; euphorbiaecola 42; euphor-
biae-pubescentis 42; herrerana 72; hibi-
sci-cannabini 60; kashotoensis 84; mi-
cheliae 62; myxa 55; oroxyli 64; panto-
leuca 42; pavettae-indicae 64, 66; peti-
la 42; phyllanthicola 89; phyllanthi-ni-
ruri 89; sapii-sebiferi 66; stillingiae
66; tectonae 68; timorensis 45; trematico-
la 89; trematis—-orientalis 45; triumfet-
tae 90; viticis 70; weberi 70
Cercosporidium bouganvilleae 98
Chaetopsina 192, 195; auburnensis
195; fulva 192; ivoriensis 195; ramifera
192; romantica 192-194
Chaetopsis 195; grisea 195
Chrysosporium 439
Ciborial 2,285) americana ole pece:
ana (43 spe | 2540 22S pen) OMmmecE anor
"sp. 1182" 2-4
Ciborinia, 1, 4, 16. 9-10;> hirsutame.
Geeta spa AGO es
Ciboriopsis 18
Cladosporium 233, 403
Coccocarpia erythroxyli 207
Cochliobolus” “353=857)5 9 359)smo7 Opmroi ae
373; australiensis 353, 373; bicolor 355;
boutelouae 355; carbonum 355; chloridis
355; ‘eymbopogonis (355, §356, "B50. oan
cynodontis 355; dactyloctenii 355, 358;
geniculatus 355, 356, 358; hawaiiensis
355, 358; heterostrophus 353, 355-358,
COs Sihln Bii4e silos
homomorphus 355; intermedius 355; kusa-
noi 355, 3583 lunatus (355,09 G06; 6 oe
melinidis 355; miakei 355; miyabeanus
355; nisikadoi 353, 373; nodulosus 355,
358; palmivora 355; peregianensis 355,
358; perotidis: 355, 356, 356: “raveneia
355, 356; sasae 355; sativus 355; setari-
ae 3552 sitharamil, 355s spicifer @eé5o.
356, 358; sporoboli 355, 358; stenospi-
lus 3553 tripogonis 355, 356; 3567) 259%
CGILICTIMGSA COs LILClLomtdemoos
Colletotrichum gloeosporioides 507
Conferticium 492, 493
Conidiobolus 284, 288-290; subg Capil-
lidium 290; subg Conidiobolus 290; subg
Delacrotxia’ 290; apiculatus 273, ~ 275,
276, 280, 282-284, 286-290; coronatus
287; destruens 273, 289, 290; major 283-
, 284, 286-288, 290; obscurus 286; papil-
latus 288, 290; pseudococci 273, 283,
286-290; thromboides 289
Coriolus 142, 143, 149; hirsutus 123,
147, 149; pubescens 143, 147, 149; versi-
color 143, 147-149; zonatus 143, 148, 149
Cornucopiella 562
Corticium 493; karstenii 565, 566; mol-
Vem565
Cortinarius 520, 521; aureifolius v
hesperius 519; fibrillosipes v_ fibrillosi-
OS DAO, 5225 salle Bile. W570, Gyr
Corynascus thermophilus 439, 440
Corynespora cassiicola 362
Crocicreas 26, 326.) 330: carpenteri
328, 329
Cryptomycina 263
Corvulartiqgeesss. 355, 359-361, 363,
365;,. 367-3733 affinis 368: andropogonis
365, 368; borreriae 365; brachyspora
368; catenulata 365, 368; clavata 368;
coicis 367; cymbopogonis 355, 367, 368;
deightonii 365; eragrostidis 368; fallax
368; geniculata 355, 365, 368; harveyi
368; inaequalis 368, 372; intermedia
355, 368; ischaemi 365, 368; lunata
Bl0e 3vo%, 303,. 305, 367, 368; oryzae
368; ovoidea 368; pallescens 368; prasa-
dii 368; protuberata 368, 369, 372; ri-
chardiae 367, 368; robusta 365; 367,
368, 372; senegalensis 368; trifolii 368;
tritici 365, 368; tuberculata 365; uncina-
ta 365, 368; verruciformis 365, 368
Custingophora 187
Cylindrocladium 265, 266, 268; citri
266; crotalariae 266; curvatum 266;
floridanum 265, 266, 268, 269, 271; ilici-
cola 266; spathiphylli 265, 267, 268-271
Cytospora 562
Cytosporinal 1231245 91260 128)" 129
Dacrymyces deliquescens 384
Dactylaria 112
Daedalea striata 351
Degelia gayana 207
Deightoniella torulosa 362
Delpinoina 262, 263
Dematium 233
Dichaena 263
Dichostereum 465
Dicranidion 112; fragile 112
Didymostilbe Boe 135% capillaceae
139; coccinea 139; coffeae 133-135; cop-
Bophilagel3s5,) 39s seichileriana 136. 135
#37, 139; ellisii 135, 137, 138; kamatii
135, 139; obovoidea 137, 139
Diplorhinotrichum 112
Drechslera 355, 360-364, 369, 372,
373; avenacea 362; biseptata 361, 372;
chloridis 355, 373; dematioidea 372; ery-
throspila 369, 372; papendorfii 361; tri-
septata 369, 372
SPT
Elaphomyces SNE: decipiens 398;
granulatus 396, 398; verrucosus 396-398
Empusa apiculata 276, 283; v apicula-
ta. 287} vi major 283, 287
Endophragmiella 152, 153; biseptata
155; canadensis 153; constricta 152-155;
corticola 155; ellisii 155; fallacia 155;
hughesii 155; hymenochaeticola 155; on-
tariensis 155; pallescens 153; tripartita
155
Entomophaga gryllii 283
Entomophthora 274, 283; apiculata
Ss llsiy PIs O83, 5, Aso conhicics
288; destruens 273, 288, 289; major
283; pseudococci 273, 286; pyralidarum
273-276, 278, 282-284, 286, 288
Entyloma 106; compositarum 106; gail-
lardiae 103, 104; gaillardianum 103,
104, 105
Epithele 461-467, 472, 479, 480, 486,
496-498; capitata 461; citrispora 463,
464, 466, 467-469, 498; dussii 461; e-
fibulata 463, 464, 466, 470-472, Mdfifs
496, 498; fulva 462; fusispora 469; gal-
zinii 462; guadeloupensis 463, 464, 467,
473-475, 496, 498; interrupta 463, 464,
466, 472, 475-477, 479; macarangae 463,
464, 466, 477, 479, 498; macrospora
461; malaiensis 463, 464, 466, 469, 479,
480, 498; nikau 462, 467, 479, 481,
482, 496, 498; ovalispora 463, 464, 467,
469, 482, (483, 2987 Step. 15697" “263,
467, 484, 485; typhae 461-464; vermi-
fera 461
Epithelopsis 462, 497; fulva 462
Erioderma neozelandica 207
Erynital "274; “2/6202, e204 i 200s eos
neopyralidarum 273, 276-278, 280-284, 287
Erysiphe 417; aquilegiae 428; asperi-
foliorum MAGS betulae 422; cicho-
racearum 424, 428; v cichoracearum
424; v poonaensis 422-424; convolvuli
428; cynoglossi 427; depressa 428; gale-
opsidis 428; graminis 428; heraclei 428;
hyperici 428; pisi 428; polygoni 428;
poonaensis 422; urticae 428
Eutypa 123, 124, 128-130; acharii
124; armeniacae 123-130; lata 124; ludi-
bunda 124
Exidia glandulosa 381, 383; nucleata
SOL 7 365
Exidiopsis effusa 382, 383; grisea 382
Exophiala 234; gougerotii 232, 235;
jeanselmei 234, 235; mansonii 235
Exserohilum 362-364; turcicum 362
Flagelloscypha kavinae WAZ, 149;
minutissima 142, 149
Fomes fomentarius 142, 143, 148; ja-
ponicus 297; lucidus 297; oerstedii 314,
318; resinaceus 308, 309; subamboinen-
sis 302
Busanium 9O93) 094s) 6403, ) 404-m sect
Liseola 393; moniliforme 394; prolifera-
tum 393, 394
578
Ganoderma 293, 296; sect Characoder-
ma 296, 318; applanatum 294, 323; ar-
gillaceum 308, 310; atkinsonii 325; biba-
diostratum 295; chaffangeonii 308, 310;
cupreum 322; fornicatum 322; laccatum
297 Vlorentzianum. 09, “Silom usi2> 31s:
322; lucidum 293=298; 300, 304, 309,
B27 TSO, wl Ose 2 Zao eoy NuLtiplicatum
VieViltali SGOL Oo een ILTG Umm oOOs eG LO
322; oerstedii 293, 296, 302, 306, 312,
iA, Sil, Bilss Sil, Seb crelouliioraiats. Sulkeie
323; -parvulum, 295; platense 9308, 310,
322; polychromum 308, 310; praelongum
308, 310; pseudoboletus 297; f montanum
325; pulverulentum 308, 310, 322; punc-—
tisporum 295; renidens 298; resinaceum
DOSS. OO, BOA, BOs, Bio, Sls sill
322; sceleton 322; sessile 294, 300, 304,
308, 310; sessiliforme 296; subamboinen-
se 304; v laevisporum 293, 295, 298,
ole, SO, BO, AIA, | SHINAR SING AT Penis. SY2e
v subamboinense 295, 304; subfornica-
tum 296; subincrustatum 308, 310, 322;
subperforatum 308, 310; sulcatum 321,
222 tuberculosum | 29385) 9c02, 306, SIZ,
Gulyé, CAMS, VAS rxoyevenabin! PASS ASI» OVA
STOR Slee lee eZ
Geopora 397, 400; cooperi f cooperi
396, 399, 400; f gilkeyae 396, 399, 400
Gigaspora 429; nigra 432, 434; reticu-
lata 429, 430, 432, 434, 435
Gloeocystidiellum 462, 492, 493, 497;
humile 493; lactescens 493; porosum 492
Glomerella cingulata 507
Grammothele 172, 173; delicatula 172;
lineata 172; macrospora 172; setulosa 172
Grammothelopsis 172
Gyromitra 165; ambigua 165; infula
165, 166; sphaerospora 165
Hartiella coccinea 139
Hebeloma crustuliniforme 515
Helminthosporium 354, 355, 359, 371;
subg Eu-Helminthosporium 353; coicis
367; geniculatum 359; homomorphus 374;
kusanoi 354; maydis 354; micropus 354;
oryzae 354; sativum 354; setariae 354;
stenospilum 354
Helvella sphaerospora 165
Henriquesia 263
Heterochaete 461, 497
Hexagonia bartlettii 174; heterospora
174
Hirneola auricula-judae 385
Humaria rhodoleuca 441, 444, 447,
449-452
Hydnum 486
Hymenoscyphus 112
Hyphodontia efibulata 473
Hypochnicium 565; analogum 571; bom-
bycinum 565-567; v bombycinum 565; v
irpicoides 568, 571; v_ pinicola 565;
cystidiatum 571; eichleri 565, 568, 569,
571; karstenii 565, 566-568, 571; punctu-
latum 565, 568-571; vellereum 571
Hypochnus 461; sect Epithele 461; dus-
sii 461
Hypogymnia 157, 158, 160, 161; dupli-
cata 158, 159; enteromorpha 157-159;
heterophylla 157, 159, 160; imshaugii
158; krogii 158; laminosorediata 161;
mollis 157, 160, 161; occidentalis 157,
158-160; tubulosa 161
Hypotrachyna sinuosa 207
Hysterangium separabile 511
Idriella 112
Incrustoporia 505
Inocybe 259; jalapensis 259; jamaicen-
sis 259; lanuginella 259; tropicalis 249,
25 i ZOO en LOO
Inonotus 116; hastifer 116
Junghuhnia 500; collabens 504; micro-
spora 503, 504, 505; nitida 505
Lactarius deceptivus 240; deliciosus
513; vellereus 240; volemus 240
Lambertella’ 1; 10)" 13.175 lS cmrcosnias
maris 10; myricae™ 105 11-5 isp ommliece
10, 12; “sp. 1959" 10, 13, 1458 -vibeerm
11; zelanica 10-12
Lamprospora Waly uA S carbonaria
118; haemastigma v gigantea 120; multi-
guttulata 121
Lanzia 2, 15, 17; “183 ‘echinophulla
Sl” “spew 187!" 21550) 16a (3 eS eecOcm
NS. IGG
Laterispora brevirama 155
Leioderma pycnophorum 207
Lepista (2535) 5 glabellay 253.
DMs, Wein PAsvAy. Pe
Lethariicola 262, 263
Leucogyrophana 565
Leucoscypha 441, 444, 446; borealis
458; leucotricha, "4442 446) sovalilane Gane
rhodoleuca 447
Libertella 562
HODaAGLa uo imc o ae Ces amplissima
224; retigera 207; scrobiculata 207
Lophophacidium 263
Lyophyllum 520, 523
singeri
Megalocystidium 492, 493, 497; lactes-
cens 492; leucoxanthum 492, 497; luri-.
dum 492
Megasporoporia 172, 173; cavernulosa
172, 174, 175; hexagonoides 172, 174,
176, 177, 180; mexicana 172, 174, 178
180; setulosa 172, 174, 180, 181
Melanconiopsis inquinans 562
Melanconium 562
Melanoleuca 254; brevipes 254; pseudo-
luscina 254; tropicalis 229, (2515992535
254; tucumanensis 254
Micraspis 263
Microdochium 112; phyllanthi 112
Microsphaera 417, 422; alni 417, 418;
astragali 428; berberidis 428; betulae
422; caprifoliacearum 422; diffusa 420;
divericata 428; euonymi 428; friesii
420; v dahurica 420, 421; v_ friesii
420; grossulariae 428; hypericacearum
428; juglandis-nigrae 418; lonicerae
422, 428; magnifica 418, 419; miurae
420, 422, 423; miyabeana 422; neglecta
417-419; nemopanthis 418; neomexicana
LUO Le ORNL Ie mornata: 416.) 2225 vy euro—
paea 422; v ornata 422; penicillata
(2S Se tolmin Al jou 4182. prasadiis 420);
ulmi 417
Moellerodiscus 2, 3, 18, 25, 28; he-
derae 19, 23, 24; iodotingens 19; subsp
canariensis 19, 22, 23; subsp iodotin-
ens 9 me20)ue22e 2058 MUSaes 223) sp.
1881" 19, 26, 27; tenuistipes 22
Monilia 28
Moniliella 234
Monilinia 28
Myceliophthora 436, 439; fergusii 440;
hinnulea 436, 438-440; thermophila 440;
vellerea 440
Mycobonia 461, 497
Mycothele 462
Myrioconium 9, 28
Nakataea 361, 367, 369;
367-369
Nannfeldtia 263
Nanoschema 562
Nectria 373
Neocosmospora 387, 394; africana 393;
striata 387, 393; tenuicristata 387, 388,
ol, 90m oon; vasinfecta 387, 393
Neottiella 441, 446, 458; aphanodicty-
on 441, 442, 458; ovilla 447; rutilans
446
Nephroma 227; australe 204; cellulo-
sum 204; kuehnemanii 228; lepido-—
phyllum 204
sigmoidea
Octavianina cigroensis 249, 257, 259,
260; laevis 260; tuberculata 260
Octospora 350; applanata 350; rubens
457
Odontotrema 262, 263
ODONTOTREMATACEAE 263
Odontura 262, 263
Oospora 233
Ophiobolus 353% 354, Si hile cariceti
353; herpotrichus 354; heterostrophus
353, 354, 375; kusanoi 354; miyabeanus
354; sativus 354; setariae 354
Orbimtay 107, 112: juteorubella, 112;
piloboloides! l07,.. 1085, 110, 1125 113;
xanthostigma 112
Pannaria fulvescens 207
Pannoparmelia angustata 204; wilson-
ii 204
Parmelia 162; amphibola 207; crambi-
diocarpa 162; discordans 162; kerguelen-
sis 162; protosulcata 162, 163; pseudo-
sulcata 162; saxatilis 162; sulcata 163,
164
Patella ovilla 447
Reliveeraye lO7e) ) 216,05 1222
198; avenosa 222
Peniophora 461
Peronospora 538, 539, see 540-556
Pestalotiopsis 371
aphthosa
+19
Peziza 350; applanata 350, 351; ara-
neosa 343-345; bolaris 351; v explanata
son; ) calopus: 30}. ovilla “441, 443.. .444,
446, 447, 449, 451, 452; patavina 450;
permuda 450
Phacidina 263
Phacidiostroma 263
Phacidium 263
Phaeoisariopsis argyreiae 35, 37, 39;
caesalpiniae 80, 84, 85
Freltinus): 114,. 116; 2142) «46 3220-245:
alni 142-144, 146, 149, 2438, 245-247:
andina, 114, 115; .hartieii 144...243,. 246,
247; hippophaécola 243- 247; igniarius
L226 dbs 1106, 242-247 =. fF a inis 9223.
nigricans. 143;,°145,. 1463 pints 123, 145%
populicola 142, 143, 145, 146: robustus
244, 246, 247; tremulae 145
Phialea calopus 28
Phialophora 233, 234; gougerotii 235
Phoma 354, 371, 403, 404, 406, 562;
americana 403, 404, 406, 412, 413; pomo-
rum 404, 406, 413; prunicola 404
Phyllactinia guttata 422, 428
Physma chilense 207
Phytophthora 99, 102, 538; bahamen-
SUSHL OZ: epistomium 102; mycopara-
sitica 102; operculata 99, 100, 102;
vesicula 99
Pilobolus 107, 110,) 112
Piricauda cochinensis 189
Placodes lucidus 297
Plasmopara 538, 539, see 540-566
Platismatia stenophylla 159
Poculump: l.uecseeZOnmnGOyn COs hinmumen 4.
25a (S05. N31 uN spueaison”. 2664 53t4, 6 306
sydowianum 28, 31
Podoscypha 473
Podosphaera clandestina 428
Polychidium contortum 207
Polymorphum 262
Polyporus cavernulosus 174-176; ja-
ponicus 297; laccatus 297; lindbladii
505; lucidus 293, 297, 322; v zonatum
SZZ3 Ee ChiZOpniluse LAZ 243-5 DLL LOM OLy,
Poria cinerascens 505; delicatula 174-
176; hexagonoides 176; linearis 174-176;
novae-zelandiae 505; setulosa 173, 180
Protomyces gaillardiae 104
Pseudobalsamia magnata 397
Pseudocercospora 66; abelmoschi 61;
alternantherae S55 39, 41; alter-
nanthericola 41; brideliae 60; brideliico-
la 58, 59; clerodendri 86; cordiicola
98; cruenta 49; daturina 41; dolichi
49; euphorbiae-piluliferae 35, 42, 45;
hibisci-cannabini 60, 61; kashotoensis
84, 87; meliicola 80, 86, 88; micheliico-
la 58, 62, 63; mungo 49; oroxyligena
58, 64, 65; pantoleuca 42, 47; pavet-
tae-indicae 58, 64, 66, 67; phyllanthi-
niruri 89, 91; polyalthiae 35, 43, 45;
stillingiae 58, 66, 67; subsessilis 89;
tectonicola 58, 68, 69; timorensis 45;
trematicola 89, 91; trematis-orientalis
45; triumfettae 90; vignae-reticulatae
49; vignigena 35, 47, 49; viticicola
580
70; viticigena 58, 68, 71; vitici-quina-
tae) 70; vitigena C5 49551
Pseudocochliobolus 353, 354, 356-359,
373, 374; australiensis 373; nisikadoi
B50, PCOCMNSTO
Pseudocyphellaria 197, 199, 207, 214,
PNG, CEP. DIR DT. Hae ubleyeshy Ghee
Wil, VOW, AOS, BB, Ble, Piles Bile, 275%
anomala 159; aurata 207; billardierii
205; carpoloma 204; colensoi 204; cori-
Masa AOl=W0Oe, BOB, lA, Wiles AUS, Lae
coronata 204; crocata 204, 207; delisea
198; episticta 203, 207; faveolata 198,
207; freycinetii 214; granulata 198,
204; homoeophylla 198, 203; hookeri
207; intricata 207; junghuhniana 199;
margaretae 197, 201, 203-205, 207, 208,
DIGS HNNZ 25-227 Se muLrrayid 197, 20k,
AOS AUS, YAO, BA, PG, ili, PAG jexxeslxe—
ta 204; psilophylla 207; pubescens 201,
POS-20S, DOs By BlOy Zig PAV, 225,
227; punctulata 199; rubella 204; rufo-—
Whereas ie, Ol, 20s, LOI, 2g ZO;
DIP Oe lOe O22 5.5 slibvariabt lis s20i/is
thouarsii 214, 220
Pseudographis 264
Pseudoperonospora 539, see 540-556
Pseudophacidium 262, 263
Pseudorhizina sphaerospora 165, 166
Pseudospiropes 191; falcatus 189-191;
nodosus 191; rousselianus 191; simplex
191
Psoroma durietzii 204; euphyllum 204;
pallidum 204
Pteridomyces 461-466, 486, 490, 492,
496-498; galzinii 463; lacteus 463, 466,
485, 486, 487, 496, 498; roseolus 463,
466, 488-490, 498; "sp. 8270" 463, 466,
490; sphaericosporus 463, 466, 490, 491,
498
Pulvinula 117, 118, 120-122; carbona-
ria v brevispora 118; constellatio 118;
convexella 117-121; discoidea 118; etiola-
ta llGaweclobitercm 117, sO eZ naema.
stigma 119; v gigantea 120; haematostig-—
ma 119; lacteoalba 119; laeterubra 117,
WAG: 2 miltina 120; mussooriensis
117, 120, 121; neotropica 119; nepalen-
Sis) 11 jon iVveoal ba, alZ0Lneori—
inauncss Mil. Tits. SOR Awe Mises Syde
WY G5, Lilet, | AILS Mees Gyoe EI ALI.
120, 121; tetraspora 118
Pustularia rhodoleuca 447; rosea 453,
456
Pythium 538
Ramonia 264
Rhinocladiella 233; mansonii 235
Rhinocladium 233
Rhinotrichum domesticum 252; lanosum
252; mexicanum 249, 250-252; subaluta-
ceum 250; subferruginosum 252
Rhizopogon colossus 513; parksii 513;
subsalmonius 513; villosulus 515
Rhodoscypha 441, 442, 444, 446; ovil-
la 441-443, 445-447, 449, 451-453, 455
Rhodotarzetta 441, 453; rosea 441,
GL2) 053) 451450
Rubetella orthocladii 184
Rutstroemia 17, 31, 33; calopus 28,
29; firma 30; setulata 33; sydowiana 31
Sarcinulella 557, 560, 562, 563; bank—
Siae 55/5) DDO sD Oo Ol
Sawadaea bicornis 428
Sclerospora 539, see 540-556
Sclerotinia 1, 34, 155; libertiana 34;
minor 152,153, 155s) scllerotionumme4
Scytinostroma 465
Scytinostromella 492
Sebacina incrustans 382, 383
Seiridium 370, 371
Skeletocutis 505; nivea 500; v diluta
505
Skeletohydnum 462-464, 467, 469, 488,
4905 297 nikaw 46658464
Skyttea 262, 263
Smittium 167, 170, 183, 186; cellaspo-
ra 183, 184, “166 -)seculicis slo/ pmo
culisetae 167, 186; longisporum 167,
169, 170; macrosporum 170; orthocladii
184; simulii 186 :
Sorosporium androsaces 104
Sphaeronaemella helvellae 165
Sphaerotheca alchemillae 427-428; a-
phanis 427; epilobii 428; ferruginea
428; fuliginea 428; macularis 428; pan-
nosa 428
Spilomela 262, 263
Sporidesmium 361, 367, 369; sclerotivo-
rum 153; tropicale 367-370
Sporothrix 234; schenkii 2335 235
Sporotrichum DBZ beurmannii PRS
234; gougerotii 232-236
Stachybotrys 187
Stenella 56; bougainvilleae 96, 97;
canthii 35, 51; cedrelae 80, 90, 93;
coffeae 58, 70, 73; cynanchi 80, 92,
94; dioscoreicola 35, 53; garugae 58,
72, 75: myxa 35, 55; oroxylicola 589
74, Tes stephaniae 35, Bays xerom-—
phigena 58, 76, 78
Stereum 461, 462; hirsutum 148
Sticta 197, 207, 216, 222; canarrensre
222, 227 “dufouril 222,. 722755 atiinows
207, 222, 224: richardii v rufoviresceme
205; weigelii 207
Stilbella 133; erythrocephala 133
Stromatothecia 262, 264
Tarzetta 456; rosea 456
Teratosperma 155; oligocladium 155;
sclerotivorum 153, 155
Thecaphora 104; androsaces 1035 SlOgg
androsacina 103, 105; jubilei 104
Thelephora 493; avellana 351; bomby-
cina 566
Thielavia heterothallica 439, 440
ToeUlawZ33
Trametes subserpens 180
Tremella 384; fimbriata 384; foliacea
384; lutescens 384; mesenterica 383, 384
Triblidium 264
Tryblidiopsis 264
Tuber 397, 400, 401; dryophilum 396,
400, 4O1; levissimum 396, 400, 401
Tubulicium 461
Tyromyces graminicola 501, 504
Uncinula 425; adunca 428; ampelopsi-
GiSm ci ID UCOLN IS | 4265.) Climtondiln "425.
427; miyabei 425; necator 427; v. ampel-
opsidis 425, 427;. cleosa 425, 427; v
oleosa 426; v zhengii 425-427; prunastri
428; subfusca 427
Ustilago androsaces 104
Vakrabeeja 361
Vararia 465; aurantiaca 473
581
Veluticeps 461, 497
Vermicularia gloeosporioides 507
Vesiculomyces 461, 465, 492, 493,
497; chelidonium 493; citrinus 492, 2493;
496-498; epitheloides 466, 485, 493-498;
humilis 493; lactescens 493, 496, 498;
leucoxanthus 493; luridus 493; sulcatus
493
Wangiella dermatitidis 234, 235
Whetzelinia 34; sclerotiorum 34
Wrightoporia 504
Xerotrema 262, 264
Xylogramma 330
582
ERRATA, VOLUME ELEVEN
Page 381, last line: for 637. read 367.
ERRATA, VOLUME FOURTEEN
14(2) cover line 27: for 1981 read 1982
ERRATA, VOLUME FIFTEEN
Page 197, add: leaves of Ammophila arenaria, 17.VII.1963,
leg. OB. no. o2iG0p. tS).
Pages 529, 530: omitted
ERRATA, VOLUME SIXTEEN
Page’ °35; ‘line 23: for S. read Stenella
80 ba: for C. read Cercoseptorta
5 Bos for S. read Selerotinta
174 44: for P. read Polyporus
176 10% for P. read Polyporus
180 18: for subsperpens read subserpens
204 Shoe for P. read Pseudocyphellarta
235 He for Sp. read Sporotrichum
28: for S. schenkit and Sp.
read Sporothrix sechenkit and Sporotrichum
234 1 for Sp. read Sporotrichum
255 oo for S. read Sporothrix
236 ac for Sp. read Sporotrichum
278 38: for EF. read Erynta
282 38: for FE. read Entomophthora
39: for EF. read Erynta
283 55 for grylly-type, F. read gryllt-type, Erynta
28498 28,5) O28 for FE. read Entomophthora
286 Bs for EF. read Entomophthora
287 38: for EF. read Erynta
293 13: for tuberculatum read tuberculosum
297 ifs for F. read Fomes
55 202 for THE PYTHIUM read THE GENUS PYTHIUM
ACJ ee 27, 55% for FE. read Eptthele
464 As for efftbulata read eftbulata
472 34: for E. read Eptthele
473 41: for Hypodontia read Hyphodontia
492 34: for Conforttetum read Conferttctum
PUBLICATION DATES FOR MYCOTAXON
Volume 15 July 15, 1962
Volume !6, No. 1 October 27, 1982
Ltt
TABLE OF CONTENTS, VOLUME SIXTEEN
No. 1 October-December 1982
A preliminary discomycete flora of Macaronesia: Part 5,
ScCrerorlupacese< A,TNDA Mos KOHN na cssle «can ore eitalcts sch 6 56 6 o aieie sie eeier 1
Studies on hyphomycetes from West Bengal, India,
I. Cereospora and allied genera of West Bengal, 1. 35
II. Cereospora and allied genera of West Bengal, 2. 58
III. Cereospora and allied genera of West Bengal, 3, JO-MIN YEN,
Pm De, (Rec LING s iors, riavese ay ctety ain Mepeteteteertenens + sic fa Ney chums init cane cpete we 80
Studies on parasitic fungi from South East Asia, 45. Parasitic fungi
HLromenalay sia, 225) .JO=MIN= YEN ~G, GLORTAVIAMS, . 54 athe a ak State as Bees 96
Phytophthora opereulata sp. nov., a new marine fungus, K. G. PEGG §&
aN GCORN woe ty as hice a ote pealgle sv 6G ine Sientee wep Ca. 8 shee whale stan emis © Bin etme ie 99
Thecaphora androsactna and Entyloma gatllardtanum, new species of
Ws me aera heS 0 KALMAN VANKY s 2c saccuatetoe © Ae eee & ote: boa eect tastes bee sde too laseas 103
A new species of Orbilta from Canada, J. H. HAINES & K. N. EGGER.... 107
Phellinus andina Plank §& Ryv. nova sp., S. PLANK §& L. RYVARDEN...... 114
A reinvestigation of the north-west Himalayan Pulvinulas, RISHI
ROME cael opens she oh eite Git 0 oss iay'aiceus roca. o's ails Laelye tel Ses ove 4. aos) sta tabial oie! MeninteneManieta “aye ele Tes sty!
Isolation and identification of Zutypa armentacae from diseased
grapevines in Washington State, D. A. GLAWE, C. B. SKOTLAND &
VM ee OLE a acehele tha a tate thre a arerech. le eee cc Oise a cereiel terme ee aa eee Bnettrabe Socetare 123
Stilbellaceous fungi 1. Ditdymosttlbe, J. L. CRANE §& ADRIANNA D.
BAe Gay poe ates Peied See Gi ws Saat Ot a aS NELa Sia, Nee aunt a ee, aim aed ats Cee ate: weno Nabene 133
Variability of spores of different basidiocarps growing on the same
mycelium in the Aphyllophorales, ERAST PARMASTO §& ILMI PARMASTO... 141
A new species of Endophragmiella from sclerotia of Selerotinta
Mee Ge UCN Dr ba. DUNN). Res rate Peak, eae ae, Sees tas £52
Three new species of Hypogymnta from western North America
(Lichenes: Hypogymniaceae), LAWRENCE H. PIKE & MASON E. HALE, JR.. 157
A new species of Parmelta (Lichenes) with protocetraric acid, MASON
EATEN ETRY OOM one ec re acs ee: ata ebare Sie a oie le tend tena, Gata Reig a siohe a mame. Ws alaeoate nee e 162
A new host for Sphaeronaemella helvellae — Pseudorhtztna
SOG FOSPONds DONALD Hy PRLOLERs fc ivan sa he Seo eine dies atte tienes es 165
Smitttum Longisporum, a new Harpellales (Trichomycetes) from
chironomid guts, MARVIN C. WILLIAMS, ROBERT W. LICHTWARDT §&
SUECHENONe “VE LERSON erat. sok sis oe ol u:g oe eae ee ate male anes apes Phare e hic etenae 167
Megasporoporta, a new genus of resupinate polypores, L. RYVARDEN,
RCM NOrMde GMs ERAS GHENDERGs <2 as atecs-a eis Sikes cr sieis nie tente nace os fale ole ees ars 172
Smtttium cellaspora, a new Harpellales (Trichomycetes) from a
enproromid nanacuc;,.MARVIN: GC" WILLIAMS 7.5455 so emiew sees ca ds ors wien 183
Notes on hyphomycetes.
XLII. New species of Acrodictys and Pseudosptropes from South
Africa. 187
XLIII. Concerning Chaetopstna romanttca, G. MORGAN-JONES.......... 192
Phycosymbiodemes in Pseudoeyphellarta in New Zealand, B. RENNER §&
Dees OALAAWAN eit 2s. 5 Shure ae late otk WS eee ewe ware Ne GFL alate aveet ne ins. es 197
A note on Sporotrtchum gougerotit Matrouchot 1910, MICHAEL R.
MEGINN TOG. ULBEROKAJELLO Osis i's shee ata stellen yee MTD ee EA clels inl eels Zoe
Paraffin embedding and semithin pene of basidiocarp tissues,
REULYS Are GRAT TO: GYDARRY Le Wes GRUND 2s ie onc we he ee ieee eae aes 259
Basidiospore variation in local populations of some Aphyllophorales,
ERAc er ARMAGE OG). LMT -PARMAGTOs © o.k't ace a/ace esse ee tate Sis eres ce aeaa ers 243
New species of fungi from the Yucatan Peninsula, GASTON GUZMAN...... 249
Two new families in the Ascomycotina, D. L. HAWKSWORTH & M. A.
DEW: ste vetesd achavele tle © aceis cio wuers sos Shere os e'< 4 ela Wee @ oe eiehe mala teauerenoname 262
Lv
Cylindrocladium spathiphylli sp. nov., CALVIN L. SCHOULTIES, NABIH
B. BL=GHOLLG SALVATORE: A ALPIERT , JR atic ucts w ss ee ee eee eee 4 ee
Erynta neopyraltdarum sp. nov. and Contdiobolus aptculatus,
pathogens of pyralid moths: components of the misdescribed species
Entomophthora pyralidarum [Zygomycetes, Entomophthorales], ISRAEL
BENS AE VEN We tubaievsd pons) «tise inane or 'astalatnrean ances & ee cate ie a. Seer ee teen ett ee Zia
Survey of the Argentine species of the Ganoderma luctdum complex,
MARTA. EB. BAZZALO” GORGE WE. WRIGHT. ..0.c., erase yan cas | oben Cone 293
Two new species of Helotiales from the eastern Himalayas, R. SHARMA
GS RIGHARD PAP ROR at tler te g's a ote, tng eimnel ot Ak ars lang ee tie cee ene eee 326
Revue des” Livres iG tLe HENNEBERT isa se ede masa ertee eo bs iene eee 331
John S. KARLING; E.J.H. CORNER; Walter JULICH; H. JAHN; H.-U. KUCK;
S.T. CHANG & W.A. HAEYES; A.J. VAN DER PLAATS-NITERINK; David H.
GRIFFIN; J. BREITENBACH & F. KRANZLIN; James B. SINCLAIR; Rivka
BARKAI-GOLAN
Notices:
International Mycological Association Record of Business Meeting.. 335
Further suggestions itor MYCOTAXON, authors. 0 awak ce oe eee ee 340
No.2) January-March 1983
Sanctioned epithets, sanctioned names, and cardinal principles in
's. Pers "wands ev Fr 4? seatations ) ORICHARD? Pa! (KORE. . si let. «5 eeeae eee 341
On the genera Cochltobolus and Pseudocochitobolus, J. L.-ALCORN..... 353
Tremelladiesdot [sraehyiNe oBINYAMING. ac . sees es a's cee eles Gite ee ie ee 380
A new Japanese species of Neocosmospora from marine sludges, SEIICHI
UEDA.“@ SHUNATCHT WDAGAWA Se sros.0 22% cacalete ass ieiats olvleteleperers Gitshe 2 (etaite see Rees 387
New records of hypogeous Ascomycetes in Arizona, JACK S. STATES..... 396
Studies in the genus Phoma. 1. Phoma americana sp. nov., G.
MORGAN=JONESMG. JAMES@). © WHITE s,s ya elo « > © eee 6 ate ip ae es see 403
A new ispecies ot: Amantta,, DAVID Ti .JENKINS&2) 2°. ..99 segs es ee eee 414
Descriptions of new species and combinations in Microsphaera and
Rrystone “ChPT ig, UWEUBRAUNG SS as cansd tee ge nerete cere alec eachom ee eee eee 417
Taxonomic notes on some powdery mildews (II), UWE BRAUN............. 425
Gigaspora rettculata: a newly described endomycorrhizal fungus from
New England, R. E. KOSKE, DIANE DOUD MILLER §& CHRISTOPHER WALKER.. 429
A new thermophilic species of Myceltophthora, TAKEYOSHI AWAO &
SHUN= TCHLSUDAGAWA 6. « ceiteowia ious avrete. tals ie Riles dain a"aie. Sunde teieta, ob xarel tre, ehole ees bel eueiaeme 436
Operculate Discomycetes from Rana (Norway) 5. Rhodoscypha gen. nov.
and Rhodotarzetta gen. nov., HENRY DISSING §& SIGMUND SIVERTSEN.... 441
Basidiomycétes Aphyllophoralés epitheloides étalés, J. BOIDIN § P.
LANQUET UNG on 2 fc eee fi, teal hank NIG Bre cet ONO en ae 461
New South American resupinate polypores, MARIO RAJCHENBERG.......... 500
Colletotrichum gloeosportotdes (Penzig) Penzig et Saccardo, S. R.
PENNY COOK: it suete ae oat folk e's ala) gi nuetau nls ale mea! aie Case ngeh ate aie Ore hate Reema 507
Ectomycorrhizae of selected conifers growing in sites which support
dense growth of bracken fern, JAN ACSAI & DAVID L. LARGENT......... 509
Mycorrhizae of Arbutts menztesit Pursh. and Arctostaphylos manzantta
Parry in northern, California, JAN-ACSAI § DAVID-L.. LARGENT...-2en. 519
Check-list of Romanian Peronosporales, 0. CONSTANTINESCU & G.
NEGREAIN sisi hipaa tote iets Ao 0S.) o-lalislsg- 0 ool» 9a le ied Sateen’ va eae eae Meee eau ai eee Son
Sarctnulella bankstae gen. et sp. nov., a Coelomycete with a unique
method of conidial dispersal, B. C. SUTTON & J. L. ALCORN......... 557
Cultural studies in Hypochntcium (Corticiaceae, Basidiomycetes),
NIELS. HALLENBERGs onset aisoetate. aleve coved alo. pode Gye oi atk etaie ote el gt miele |e 2 Se 565
NOS ron ADL ANS aPOr aM), eras sd. gn ge teraaitsa ate Cee aos ela toe ee 572
PONE es Ne ccm tale als! vista ale civigl o Ae Oe Se RM ee Me na iced Mel dt ee ee ae)
VED OmrunZOus “anda lichen taka, sack ste ane 8 Ua cis Met ee a eee 576
DSRS yO MAS TO PINES ERC UAB See Aaa ee RS ane en Te SURG 582
MyLOUAXGN publication dates, 15 and 1601) gees oe. Oh). oe 582
ERIS L008 gee a i en ar re ane ge eke ed a a re Oe pa vt
vt
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als
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